The effects of abrupt topography on plankton dynamics

Plankton population dynamics in the upper layer of the ocean depends on upwelling processes that bring nutrients from deeper waters. In turn, these depend on the structure of the vertical velocity field. In coastal areas and in oceanic regions characterized by the presence of strong submarine topographic features, the variable bottom topography induces significant effects on vertical velocities and upwelling/downwelling patterns. As a consequence, large plankton and fish abundances are frequently observed above seamounts, canyons and steep continental shelves. In this work, the dynamics of an NPZ (nutrient-phytoplankton-zooplankton) system is numerically studied by coupling the ecosystem model with a quasi two-dimensional (2D) fluid model with topography. At variance with classical 2D approaches, this formulation allows for an explicit expression of the vertical motions produced when fluid columns are squeezed and stretched as they experience changes of depth. Thus, input or output of nutrients at the surface are associated with fluid motion over the bottom topography. We examine the dynamics of a cyclonic vortex over two basic topographies: a steep escarpment and a submarine mountain. We show that plankton abundance over the escarpment is modulated by the passing of topographic Rossby waves, generated by the vortex-topography interaction. In such configuration, advection effects driven by the flow over the escarpment are of limited relevance for the dynamics of biological fields. By contrast, we find that the flow resulting from the interaction of a vortex with a seamount is sufficiently strong and persistent to allow for a remarkable increase of nutrients, and a corresponding enhancement of phytoplankton and zooplankton concentrations. Over the seamount, advection effects associated with trapped flow perturbations around the summit play an essential role.     

The landward distribution of oceanic plankton and micronekton over the west Florida continental shelf as related to their vertical distribution

The landward distributions of 69 plankton and 92 micronektonspecies over the west Florida continental shelf were examinedin relation to their vertical distribution in the eastern Gulfof Mexico. Using linear and power-curve regressions, it wasfound that extent of landward occurrence is significantly correlatedwith bottom topography in terms of bottom depth and distancefrom the open Gulf. Epipelagic plankton species were distributedconsiderable distances across the shelf whereas the mesopdagicshrimp and fish species were not found landward of slope stations.Possible factors affecting landward distribution, such as currents,vertical migration patterns, and predation, are discussed.     

地形异质性对云南普洱季风常绿阔叶林物种多样性的影响

地形异质性通过调控树木生长所需的养分、水分和光照等而成为亚热带森林结构与物种组成的重要驱动因子。但是, 地形异质性对季风常绿阔叶林物种多样性及其分布影响的研究还相对较少。该文基于云南普洱30 hm²森林动态监测样地(大样地) 750个20 m × 20 m的样方调查数据, 以海拔、坡度、凹凸度和坡向4个地形因子为变量, 采用C均值模糊聚类分析大样地的地形类型, 进而分析不同地形条件下的群落物种组成及群落物种多样性; 采用Torus转换检验法, 探讨物种与地形关联性, 为季风常绿阔叶林生物多样性保护提供科学依据。研究结果表明, 大样地可分为山脊、陡坡、缓坡、高谷和沟谷等5类地形, 地形面积分别是8.00、6.04、7.68、2.76和5.52 hm²。大样地中胸径(DBH) ≥ 1 cm的木本植物个体153 418株, 分属79科179属271种。5类地形中, 物种丰富度、不同径级的植株密度和比例明显不同, 多样性及优势物种多度分布具有较大差异。种-面积曲线表明, 同等面积条件下, 随着取样面积增加, 山脊的物种丰富度始终最小, 高谷次之, 沟谷的物种丰富度始终最大。种-个体数累积曲线表明, 随着个体数增加, 山脊物种丰富度的累积速率最小, 种丰富度增加缓慢, 高谷次之。在被检验的123个物种中, 与地形相关的物种有83个, 高达67.5%的物种与至少一类地形存在显著相关关系。山脊和缓坡中与地形具有显著负相关关系的物种数超过显著正相关的物种数; 而与陡坡、高谷和沟谷显著正相关的物种数高于显著负相关的物种数。普洱大样地地形异质性对物种多样性维持的贡献率为7.8%。     

Simultaneous measurements of small-scale physical dynamics and zooplankton distributions

Haury et al., (Deep Sea Res., 37, 447–461, 1990) describedhow turbulence and vertical shear generated by the passage ofa storm affected the vertical distributions and community structureof plankton in Monterey Bay, CA. The data presented came froma larger set of five plankton recorder collections of zooplankton,taken together with temperature, salinity, turbulent kineticenergy dissipation rate, and vertical current shear data onthe research submarine Dolphin. Here we report on the analysisof the whole data set, which represents a wide range of physicaldynamic conditions. While the overall community structure remainedrelatively constant over the sampling period, the relationshipsbetween taxonomic categories and between categories and thephysical dynamics were complex and variable; factors relatedto depth were the most important. During periods of weak winds(low kinetic energy conditions) some of the biological variabilityappeared to be related to the level of turbulent dissipation.Other relationships were not detected due to the limitationsof the plankton sampler and its interactions with the highlyvariable biological and physical conditions. In view of thesedifficulties, we recommend that future field investigationsof the relationships between physical parameters and zooplanktondistributions make greater use of acoustic and other remote-sensingtechnologies, preferably in combination with numerical simulationstudies to help guide and interpret the field work.     

Biological activity in the wake of an island close to a coastal upwelling

Hydrodynamic forcing plays an important role in shaping the dynamics of marine organisms, in particular of plankton. In this work we study the planktonic biological activity in the wake of an island which is close to an upwelling region. Our research is based on numerical analysis of a kinematic flow mimicking the hydrodynamics in the wake, coupled to a three-component plankton model.We use parameter values relevant for the Canary wake, and the main results for a realistic range of parameters in this area are as follows: (a) Primary production is enhanced in the region of the wake opposite to the upwelling zone. (b) There is a strong dependence of the productivity on the inflow conditions of biological material entering the wake transported by the main current. Finally (c) we show that under certain conditions the interplay between wake structures and biological growth leads to plankton blooms inside mesoscale hydrodynamic vortices that act as incubators of primary production.     

Roughness-dependent removal of settled spores of the green alga Ulva (syn. Enteromorpha) exposed to hydrodynamic forces from a water jet

Topographic features change the hydrodynamic regime over surfaces subjected to flow. Hydrodynamic microenvironments around topographic structures may have consequences for recruitment and removal of propagules of marine benthic organisms. The settlement and adhesion of zoospores from the green alga Ulva linza (syn. Enteromorpha linza) to defined topographies was investigated. A range of topographic size scales (Rz: 25-100 microm) was manufactured from plankton nets, creating patterns with ridges and depressions. The topographic scales span a roughness similar to that of natural substrata and antifouling coatings. Spores were removed from the surfaces by a calibrated water jet. Fewer spores were removed from the smallest topographic structure tested (Rz: 25 microm) compared to both the smooth (Rz: 1) and the roughest (Rz: 100 microm) structures. Zoospores that settled in depressions were less likely to be removed compared to spores on the ridges. The results in terms of the interaction between surface topography and hydrodynamic forces have implications for both natural substrata exposed to wave action and antifouling surfaces on ships' hulls. The possible effects of topography on increasing zoospore adhesion and offering a refuge from hydrodynamic forces are discussed.     

The influence of weather conditions (temperature and wind) on cyanobacterial bloom development in the Gulf of Finland (Baltic Sea)

Cyanobacterial blooms are common in the Baltic Sea. They are dominated by Aphanizomenon flos-aquae and Nodularia spumigena and take place in July–August. Investigations of bloom development using different approaches have been carried out in the Gulf of Finland during recent years. The ship-of-opportunity technique allows to observe the upper layer dynamics from meso- to basin-wide scale with high temporal and spatial frequency at low cost. Unattended measurements on board a commercial ferry along a transect between Tallinn and Helsinki have been conducted for 3 years (1997–1999). The influence of weather conditions—temperature and wind—on the cyanobacterial bloom development was investigated. The formation of cyanobacterial blooms was favoured by warm and calm weather, while in cold and windy conditions other species formed mass occurrences. Water temperature has been found to be the main factor controlling the initiation of the bloom, in general, while vertical stratification appeared to be the critical factor determining the intensity of the bloom at species level. The spatial distribution of the cyanobacterial bloom was determined rather by the wind-forced advection than by the possible vertical transport of nutrients in the areas of the observed upwelling events.     

Plankton cycles disguised by turbulent advection

Mathematical models used to represent plankton dynamics often display limit-cycle behavior in a range of realistic parameter values. However, experimental data do not show evidence of plankton oscillations besides externally driven seasonal blooms, casting doubts on the validity of the models themselves. In this work we show that spatial-temporal variability, coupled with advection by mesoscale turbulence, can disguise limit-cycle behavior to the point that it cannot be detected in fixed-point measurements of plankton abundance. The results presented here have more general implications as they indicate that the behavior of ecosystem models in the presence of advection can be very different from that occurring for homogeneous conditions. Care should thus be exercised in drawing general conclusions from the analysis of homogeneous ecosystem models.     

Groundwater and pore water inputs to the coastal zone

Both terrestrial and marine forces drive underground fluid flows in the coastal zone. Hydraulic gradients on land result in groundwater seepage near shore and may contribute to flows further out on the shelf from confined aquifers. Marine processes such as tidal pumping and current-induced pressure gradients may induce interfacial fluid flow anywhere on the shelf where permeable sediments are present. The terrestrial and oceanic forces overlap spatially so measured fluid advection through coastal sediments may be a result of composite forcing. We thus define “submarine groundwater discharge” (SGD) as any and all flow of water on continental margins from the seabed to the coastal ocean, regardless of fluid composition or driving force. SGD is typically characterized by low specific flow rates that make detection and quantification difficult. However, because such flows occur over very large areas, the total flux is significant. Discharging fluids, whether derived from land or composed of re-circulated seawater, will react with sediment components. These reactions may increase substantially the concentrations of nutrients, carbon, and metals in the fluids. These fluids are thus a source of biogeochemically important constituents to the coastal ocean. Terrestrially-derived fluids represent a pathway for new material fluxes to the coastal zone. This may result in diffuse pollution in areas where contaminated groundwaters occur. This paper presents an historical context of SGD studies, defines the process in a form that is consistent with our current understanding of the driving forces as well as our assessment techniques, and reviews the estimated global fluxes and biogeochemical implications. We conclude that to fully characterize marine geochemical budgets, one must give due consideration to SGD. New methodologies, technologies, and modeling approaches are required to discriminate among the various forces that drive SGD and to evaluate these fluxes more precisely.     

Characteristics of secondary flow in steady and pulsatile flows through a symmetrical bifurcation

Steady and pulsatile flow in a glass model simulating an arterial bifurcation was investigated by flow visualization techniques. Secondary flow generated at the bifurcation has a similar pattern to a vortex, called the horseshoe vortex, produced around a wall-based protuberance in a circular tube. The same flow disturbance was clearly observed during the decelerating phase of pulsatile flow. The vortex produces a stagnation point on the top and bottom wall just upstream from the bifurcation apex. When aluminium dust was suspended in the test fluid perfusing the blood vessel model, particles deposited over an area spreading from the stagnation point to the lateral corners of the bifurcation. Comparison between the present results and topographical patterns of atherosclerosis reported in the literature suggests that it is in such low shear regions that lipid deposition tends to occur most.     

Computer modeling of fluid dynamics related to a myocardial bridge in a coronary artery

Fluid mechanics associated with blood flows induced by the so-called myocardial bridge (MB) has been studied systematically using a computational fluid dynamic modeling of the Newtonian, incompressible, two-dimensional, unsteady flow in a channel with a time-dependently flushing in/out indentation. During each cycle, a train of vortex wave flow was observed downstream of the phasic stenosis and both upper and lower walls suffer severely from consistently high, oscillating wall shear stresses (WSS). Extensive studies were conducted on the influence of the Reynolds number, the geometry and the Strouhal number of the MB movement on the nature of the vortex flow and the time-dependent wall shear stress distribution. Special attention was drawn to the relationship between the vortex wave and the pressure distribution. It was found that the pressure gradient changed markedly during one cycle, which was apparently dominated by the dynamics of the indentation. A steep, adverse pressure gradient was observed when the indentation was flushing out, which corresponded to the existence of the most developing vortices. It implies the possibility that the MB in a coronary artery can produce an extremely low pressure region immediately downstream of the phasic stenosis, where elastic choking or collapse of the coronary artery might occur.     

Nutrient dynamics and pelagic food web interactions in oligotrophic and eutrophic environments: an overview

The concept of limiting nutrients is a cornerstone of theories concerning the control of production, structure and dynamics of freshwater and marine plankton. The current dogma is that nitrogen is limiting in most marine environments while freshwater ecosystems are mostly phosphorus-limited, although evidence of phytoplankton limitation by either N or P has been found in both environments.However, the same considerations apply to the availability of phosphorus in freshwater as to nitrogen in oceans. In resource-limited environments the plankton dynamics depend mostly on the internal mechanisms which act to recycle the limiting nutrient many times over within the surface waters. As the overall productivity increases, this dependence on nutrient regeneration decreases.The relationship between the stock of limiting nutrient, rates of supply and plankton dynamics must therefore be seen in the light of the processes operating within the entire food chain over quite different time scales. There is strong evidence that process-rates are mostly size-dependent and that food web interactions at the microbial level (picophytoplankton, bacteria, microheterotrophs) strongly effect the production of carbon and the regeneration of nutrients in the pelagic zone.     

Is There a Seamount Effect on Microbial Community Structure and Biomass? The Case Study of Seine and Sedlo Seamounts (Northeast Atlantic)

Seamounts are considered to be "hotspots" of marine life but, their role in oceans primary productivity is still under discussion. We have studied the microbial community structure and biomass of the epipelagic zone (0-150 m) at two northeast Atlantic seamounts (Seine and Sedlo) and compared those with the surrounding ocean. Results from two cruises to Sedlo and three to Seine are presented. Main results show large temporal and spatial microbial community variability on both seamounts. Both Seine and Sedlo heterotrophic community (abundance and biomass) dominate during winter and summer months, representing 75% (Sedlo, July) to 86% (Seine, November) of the total plankton biomass. In Seine, during springtime the contribution to total plankton biomass is similar (47% autotrophic and 53% heterotrophic). Both seamounts present an autotrophic community structure dominated by small cells (nano and picophytoplankton). It is also during spring that a relatively important contribution (26%) of large cells to total autotrophic biomass is found. In some cases, a "seamount effect" is observed on Seine and Sedlo microbial community structure and biomass. In Seine this is only observed during spring through enhancement of large autotrophic cells at the summit and seamount stations. In Sedlo, and despite the observed low biomasses, some clear peaks of picoplankton at the summit or at stations within the seamount area are also observed during summer. Our results suggest that the dominance of heterotrophs is presumably related to the trapping effect of organic matter by seamounts. Nevertheless, the complex circulation around both seamounts with the presence of different sources of mesoscale variability (e.g. presence of meddies, intrusion of African upwelling water) may have contributed to the different patterns of distribution, abundances and also changes observed in the microbial community.     

Determinants of Leaf Litter Nutrient Cycling in a Tropical Rain Forest: Soil Fertility Versus Topography

We investigated the influence of landscape-level variation in soil fertility and topographic position on leaf litter nutrient dynamics in a tropical rain forest in Costa Rica. We sampled across the three main edaphic conditions (ultisol slope, ultisol plateau, and inceptisol) to determine the effect of soil nutrients on leaf litter nutrient concentrations while controlling for topography, and to examine topographic effects while controlling for soil nutrients. Both leaf litter macronutrient [phosphorus (P), nitrogen (N), sulfur (S), calcium (Ca), potassium (K), magnesium (Mg)] and micronutrient concentrations were quantified throughout a 4-year period. Leaf litter [P], [N] and [K] varied significantly among soil types. The variation in [P], [N], and [K] was explained by soil fertility alone. Leaf litter [S], [Ca], and [Mg] did not vary among the three soil types. Macronutrient (P, K, Mg, S, Ca) concentrations in the leaf litter were much less variable than those of Fe and Al. Lower variability in essential plant nutrients suggests a great deal of plant control over the amount of nutrients resorbed before senescense. Leaf litter macronutrient concentrations varied significantly over the 4-year period, but the temporal variation did not differ among the three edaphic types as anticipated. Hence, although the magnitude of nutrient fluxes may be controlled by local factors such as soil fertility, temporal patterns are likely regulated by a common environmental variable such as precipitation or temperature.     

Seasonal plankton dynamics along a cross-shelf gradient

Much interest has recently been devoted to reconstructing the dynamic structure of ecological systems on the basis of time-series data. Using 10 years of monthly data on phyto- and zooplankton abundance from the Bay of Biscay (coastal to shelf-break sites), we demonstrate that the interaction between these two plankton components is approximately linear, whereas the effects of environmental factors (nutrients, temperature, upwelling and photoperiod) on these two plankton population growth rates are nonlinear. With the inclusion of the environmental factors, the main observed seasonal and inter-annual dynamic patterns within the studied plankton assemblage also indicate the prevalence of bottom-up regulatory control.     

Structural complexity in mussel beds: the fractal geometry of surface topography

Seafloor topographic complexity is ecologically important because it provides habitat structure and alters boundary-layer flow over the bottom. Despite its importance, there is little agreement on how to define and measure surface complexity. The purpose of this investigation was to utilize fractal geometry of vertical cross-section profiles to characterize the surface topography of the soft-bottom mussel bed (Mytilus edulis L.) at Bob's Cove, ME, USA. Mussels there have been shown previously to have spatially ordered fractal characteristics in the horizontal plane. Two hypotheses were tested. The first was that the bed surface is fractal over the spatial scale of 1.44-200 mm, with fractal dimension less than or equal to 1.26, the value for the Koch curve, our model for bed profiles. The second was that bed surface topography (i.e., in vertical profile) is less complex than the mussel bed spatial pattern (i.e., aerial view in the horizontal plane). Both hypotheses were supported. Cross-sections of plaster casts of the bed produced 88 surface profiles, all of which were fractal over the entire spatial scale of more two orders of magnitude employed in the analysis. Fractal dimension values (D) for individual profiles ranged from 1.031 to 1.310. Fractal dimensions of entire casts ranged up to mean (1.242+/-0.046) and median (1.251) values similar to 1.26, the theoretical value of the Koch curve. The bed surface was less complex than the bed spatial pattern because every profile had D<1.36, the smallest value previously obtained from aerial views of the bed. The investigation demonstrated for the first time that surface topography of a soft-bottom mussel bed was fractal at a spatial scale relevant to hydrodynamic processes and habitat structure important for benthic organisms. The technique of using cross-section profiles from casts of the bed surface avoided possible underestimates of fractal dimension that can result from other profiling methods reported in the literature. The results demonstrate that fractal dimension can be useful in the analysis of habitat space and water flow over any irregular seafloor surface because it incorporates the size, shape, and scale of roughness elements into a simple, numerical metric.     

Temporal organization of phytoplankton communities linked to physical forcing

The performance of individual phytoplankton species is strongly governed by the thermal stratification’s impact on vertical mixing within the water column, which alters the position of phytoplankton relative to nutrients and light. The present study documents shifts in phytoplankton structure and vertical positioning that have accompanied intensified long-term stratification in a natural ecosystem. Ordination analysis is used to extract gradients in phytoplankton composition in Lake Tahoe, an extremely nutrient-poor lake, over a 23-year period of records. Community structure in the 1980s was associated most strongly with resource availability (low nitrogen to phosphorus ratios, deeper euphotic zone depth), while intensified stratification dominated the phytoplankton structure since the late 1990s. Within diatoms, small-sized cells increased with reduced mixing, suggesting that suppressed turbulence provides them with a competitive advantage over large-sized cells. Among the morphologically diverse chlorophytes, filamentous and coenobial forms were favored under intensified stratification. The selection for small-sized diatoms is accompanied by a shoaling trend in their vertical position in the water column. In contrast, the motile flagellates displayed a deeper vertical positioning in recent years, indicating that optimal growth conditions shifted likely due to reduced upwelling of nutrients. As the thermal stratification of lakes and oceans is strongly linked to climate variables, the present study confirms that climate warming will alter phytoplankton structure and dynamics largely through effects on nutrient availability and sinking velocities. Intensified stratification should favor the expansion of small-sized species and species with the capability of buoyancy regulation, which may alter primary productivity, nutrient recycling, and higher trophic productivity. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Wake structure and hydrodynamic performance of flapping foils mimicking fish fin kinematics

Numerical simulations are used to investigate the wake structure and hydrodynamic performance of bionic flapping foils. The study is motivated by the quest to understand the fluid dynamics of fish fins and use it in the underwater propulsion. The simulations employ an immersed boundary method that makes it possible to simulate flows with complex moving boundaries on fixed Cartesian grids. A detailed analysis of the vortex topology shows that the wake of flapping foils is dominated by two sets of complex shaped vortex rings that convect at oblique angles to the wake centerline. The wake of these flapping foils is characterized by two oblique jets. Simulations are also used to examine the wake vortex and hydrodynamic performance over a range of Strouhal numbers and maximum pitch angles and the connection between the foil kinematics, vortex dynamics and force production is discussed. The results show that the variety law of the hydrodynamic performance with kinematic parameters strongly depends on the flow dynamics underlying the force production, including the orientation, interconnection and dissipation rate of the vortex rings.