Fine-scale distribution of sardine (Sardina pilchardus) eggs and adults during a spawning event

High-resolution acoustic and ichtyoplankton sampling with a ‘continuous under-way fish egg sampler (CUFES)’ was performed in two regions of approximately 100 square nautical miles off southern Iberian Peninsula, with the aim of studying the small scale distribution of sardine (Sardina pilchardus) adults and eggs during a spawning event. Very dense patches (246 eggs m^− 3) of recently spawned eggs with dimensions (up to 3 nautical miles wide) significantly larger than daytime sardine schools were present in both regions. Egg staging and ageing showed very little intra-sample variation, indicating a synchronous spawning period at dusk. The internal structure of the patches evaluated by variography showed very low internal variability, as if they consisted of a single unit. This hypothesis is confirmed by the acoustic finding of large sardine shoals with similar dimensions to those of the patches after sunset and throughout the night. During that period, adults were found near or in contact with the bottom, suggesting that spawning occurred at depth. A distinct patch of older eggs was found in both areas, but with a few nautical miles of horizontal separation. Their characteristics (a larger area, lower egg densities and a more irregular shape) indicate that these patches were exposed to dispersion and ‘stirring’ by physical forces, reshaping their initial appearance, while mesoscale water circulation could have displaced the core of the patches away by several kilometres within a day.     

Contribution of breaking internal waves to structure formation,energy dissipation,and vertical diffusion in the ocean

The relationship between the scale of the spectral minimum of the first differences in temperature fluctuationsL and the local value of the Väisälä-Brunt frequencyN has been analysed using the results of more than 600 soundings made in various regions of the world's oceans. Allowing for the series of theoretical and experimental indications of the fact that the vertical scaleL at the boundary between the fine structure and microstructure in the ocean exists under the effect of processes of breaking of internal waves and hydrodynamic instability with the formation of turbulent patches, and using the energy relationships, the relationship for a solitary patch has been derived through the determining parameters. Based on two expressions forL, derived experimentally and using the energy estimations, the relationships for the averaged rate of absorption of the wave field energy by patches and the coefficient of the density vertical diffusion are derived, which do not contradict a series of independent estimations fromin situ measurements.Translated by Mikhail M. Trufanov.     

Neocalanus flemingeri, a new species of Calanidae (Copepoda: Calanoida) from the subarctic Pacific Ocean, with a comparative redescription of Neocalanus plumchrus (Marukawa) 1921

Copepodites of Neocalanus plumchrus dominate the macrozooplankton biomass of the subarctic Pacific during spring. Living specimens have patches of bright red-orange pigment. During May of 1984 it was found sharing dominance with another, related species with differently placed patches of more distinctly red pigment. This is an undescribed species, which is designated here as Neocalanus flemingeri sp. nov. In the Gulf of Alaska it is smaller than N. plumchrus, but in the Bering Sea where food supplies are greater it reaches dimensions as large or larger. The morphologies of the two species are distinct, and previous confusion in the taxonomy of these forms was caused by differences in their life histories.     

Abundance, distribution and patch formation of zooplankton

The goal of studies described here was to determine the responses of zooplankton taxa to phytoplankton patches which develop in and near intrusions of cold, nutrient-rich Gulf Stream water. To achieve this goal we determined the horizontal and vertical distributions of abundant mesozooplankton taxa on the south-eastern continental shelf of the USA between 29°30′ and 31°N. The study period was from June 23 to August 16, 1981. Highest concentrations of zooplankton usually occurred in and near patches of phytoplankton. Increased phytoplankton appeared to trigger the formation of patches of the calanoid copepod Temora turbinata and the cyclopoid copepods Oithona spp. and Oncaea spp. The patches of zooplankton had greater alongshore than cross-shelf dimensions. T. turbinata responded rapidly to increased concentrations of phytoplankton by reproducing and aggregating in and above intruded waters. Oithonidae which were often, but not always, abundant in phytoplankton patches eventually attained high concentrations over most of the middle and part of the inner shelf. Their concentration and that of Oncaeidae increased steadily. Oncaeidae were not abundant in recently upwelled waters, as was T. turbinata but reached high concentrations in older intrusions when the abundance of T. turbinata remained level or decreased slowly. Both cyclopoid taxa are thought to reproduce slowly (egg sacs) compared to T. turbinata. Another taxon, the doliolids, became abundant far more rapidly in intruded waters (by asexual reproduction) than did the other three taxa. Doliolids were the most opportunistic intrusion zooplankton form. They do not regularly occur in low abundance on the shelf, as do the three copepod taxa, but develop in pulses in regions where T. turbinata and Oncaea are not abundant. Of the four taxa studied the abundance of doliolids increased and decreased most rapidly, whereas Oithona and Oncaea increased slowly and did not decrease during the study period. T. turbinata and Oncaea were most abundant at 60% of all stations in the intruding water. Doliolids and Oithona on the other hand, were mostly in the thermocline and intrusion. Whereas phytoplankton patches, which developed in intrusions, were physically induced (PAFFENHÖFER and LEE, 1988), patches of zooplankton were biologically induced.     

Modelling formation of complex topography by the seagrass Posidonia oceanica

Posidonia oceanica is a slow growing seagrass species that extends via growing rhizomes that grow only centimetres both horizontally and vertically each year. Posidonia oceanica forms topographically complex biogenic reefs of dead rhizome and sediments that are up to 4 m in height that are called “matte”. This study investigates the role of slow horizontal and vertical growth of rhizomes in the formation of topographic complexity in P. oceanica matte using agent-based modelling. The simulated infilling of landscapes by P. oceanica was run over 600 iterations (years) for 10 random starts of 150 agents each. Initial infilling rates were very slow and P. oceanica had limited cover after a century of growth. Growth accelerated after 100 years but plateaued after 400 years such that after 600 years only two-thirds of the landscape was occupied by P. oceanica. The pattern of spread of agents was initially random in direction but after larger patches were formed spread was radial from these patches. The seagrass landscape was initially highly fragmented with many small separate patches made up of a few agents each, with a Landscape Division index close to 1. Between 300 and 600 years Landscape Division declined sharply to 0.42, indicating patches had coalesced into larger more continuous meadows forming a less fragmented landscape. Perimeter to area ratio of seagrass patches declined exponentially from >1 to approximately 0.2 over 600 years of simulation. The matte developed from growth of patches and its greatest height occurred in more continuously occupied cells of the grid. The topography of the reef that occupied two-thirds of the landscape after six centuries of growth could be described as a pattern of channels between reef plateaus elevated 1–2 m above channels. These results demonstrate that development in P. oceanica meadows of three-dimensional structure, in the formation of biogenic reefs, can be explained by, and is an emergent property of, slow horizontal and vertical rhizome growth rates combined with the time it takes for the accumulation of rhizomes in any region of the landscape. As such, the model provides a parsimonious explanation for the development of complex matte topography.     

Rotational instability of a horizontal shear flow in a stratified rotating fluid

The rotational instability of a thermally stratified, viscous, conducting, rotating fluid is investigated by means of linearized perturbation equations. It is assumed that the basic horizontal flow is vertically uniform and that the horizontal shear is confined in a thin layer. By solving a simplified boundary value problem as a model of rotational instability in the sea, we have shown that the vertical wave length of the neutral disturbance is of order 10 times as large as the laminar Ekman layer thickness, and that this scale is proportional to (L/N)^1/3, whereL is the width of the shear layer andN is the Brunt-Vaisala frequency.     

Current Dynamics of Estuarine Circulation in the Lateral Boundary Layer

High spatial resolution measurements of current velocity performed by the shipboard mounted Acoustic Doppler Current Profiler (ADCP) in the lateral boundary layer of the southern Gulf of Finland during two 5-day periods are described and analysed with a focus on the dominant dynamics. The measurement site represents a small (15×20 km), relatively deep (up to 100 m) bay opened to large-scale estuarine circulation. The measurement period was characterized by calm winds and a strong seasonal pycnocline (Brunt-Väisälä frequencyN=6–9*10⁻² s⁻¹). The quasi-steady velocity field revealed polarization of currents along the shore whereas an intensive baroclinic coastal jet was observed over a cross-shore scale of 1–2 km. The level of vertical separation of the alongshore flow coincided with the pycnocline at the coast, but was shifted below it in the offshore region. The cross-shore flow was considerably weaker and showed a three-layer structure with an opposite phase between the first and second surveys. It is suggested that the observed jet resembles a non-locally forced eastward propagating coastally trapped wave. In the offshore area the alongshore flow field satisfies local geostrophic balance quite well, except in the pycnocline where strong vertical stratification exerts considerable vertical stress. As vertical velocity shear is well correlated with vertical stratification, the horizontal advection prevails over vertical mixing. Horizontal inhomogeneities of density distribution are partly explained by vertical velocities with an estimated magnitude of less than 0·6 mm/s and the spatial pattern following bottom topography.     

On the Polyps of the Common Jellyfish Aurelia aurita in Kagoshima Bay

There is the natural habitat of polyps of the common jellyfish Aurelia aurita in the Taniyama area, Kagoshima Bay. We examined the attachment substrata, density, colony structure and strobilation of the polyps. The polyps were observed only on the horizontal undersurface of floating piers. They attached specifically to Mytilus shells, solitary ascidians, calcareous polychaete tubes, muddy amphipod tubes and the gap space that fouling animals peeled off the substrata. The polyp colonies were distributed in patches. Spatial distribution patterns of the polyps within their colonies were uniform. Strobilation occurred during late December to March, when water temperatures were 16–17°C, and a large number of ephyrae were released. An increase in man-made structures such as floating piers in coastal areas may lead to bloomings of Aurelia aurita medusae. This revised version was published online in July 2006 with corrections to the Cover Date.     

Mean Vertical Distribution of the Dissipation Rate of Turbulent Energy in the Black Sea. Comparison with the Existing Models

We compare the results obtained by using theoretical and semiempirical models developed for the evaluation of the dissipation rate of turbulent energy in a stratified ocean with independent distribution of this quantity established by the authors for the active layer of the Black Sea (50–300 m) by using a one-dimensional model taking into account the balance of heat, salt, and fluid inside the layer. It is shown that, in a layer with gradual variation of the Väisälä–Brunt frequency N as a function of depth, the predominant sink of the energy of motion into dissipation N ² is ensured by the flow of energy through the spectrum of internal waves toward low frequencies and small vertical scales. On the contrary, in layers with abrupt drops of density as a function of depth (layers with jumps of density), an important role is played by the interface-type waves and the dependence of on N transforms into N .     

Present-day stress and pore pressure fields in the Cuu Long and Nam Con Son Basins, offshore Vietnam

Knowledge of the in situ, or contemporary stress field is vital for planning optimum orientations of deviated and horizontal wells, reservoir characterization and a better understanding of geodynamic processes and their effects on basin evolution.This study provides the first documented analysis of in situ stress and pore pressure fields in the sedimentary formations of the Cuu Long and Nam Con Son Basins, offshore Vietnam, based on data from petroleum exploration and production wells.In the Cuu Long Basin, the maximum horizontal stress is mainly oriented in NNW–SSE to N–S in the northern part and central high. In the Nam Con Son Basin, the maximum horizontal stress is mainly oriented in NE–SW in the northern part and to N–S in the central part of the basin.The magnitude of the vertical stress has a gradient of approximately 22.2 MPa/km at 3500 m depth. Minimum horizontal stress magnitude is approximately 61% of the vertical stress magnitude in normally pressured sequences.The effect of pore pressure change on horizontal stress magnitudes was estimated from pore pressure and fracture tests data in depleted zone caused by fluid production, and an average pore pressure–stress coupling ratio (ΔS_h/ΔP_p) obtained was 0.66. The minimum horizontal stress magnitude approaches the vertical stress magnitude in overpressured zones of the Nam Con Son Basin, suggesting that an isotropic or strike-slip faulting stress regime may exist in the deeper overpressured sequences.     

Simulation of nonlinear wave run-up with a high-order Boussinesq model

This paper considers the numerical simulation of nonlinear wave run-up within a highly accurate Boussinesq-type model. Moving wet–dry boundary algorithms based on so-called extrapolating boundary techniques are utilized, and a new variant of this approach is proposed in two horizontal dimensions. As validation, computed results involving the nonlinear run-up of periodic as well as transient waves on a sloping beach are considered in a single horizontal dimension, demonstrating excellent agreement with analytical solutions for both the free surface and horizontal velocity. In two horizontal dimensions cases involving long wave resonance in a parabolic basin, solitary wave evolution in a triangular channel, and solitary wave run-up on a circular conical island are considered. In each case the computed results compare well against available analytical solutions or experimental measurements. The ability to accurately simulate a moving wet–dry boundary is of considerable practical importance within coastal engineering, and the extension described in this work significantly improves the nearshore versatility of the present high-order Boussinesq approach.     

Dynamical structure of vertically two-dimensional estuary in steady state

The dynamical structure of a two-dimensional (depth and axial directions) estuary is studied analytically. A set of governing equations describing the time-averaged velocity and salinity in the estuary is used, where all of the external parameters (depth, width, freshwater discharge and horizontal and vertical coefficients of eddy viscosity and eddy diffusivity) are assumed to be constant along the estuary.Two dynamical relations are taken into consideration in the theory. One of them is the dependence of the longitudinal scaleL _d on the balance of longitudinal salt transport, and the other is the relation between the vertical stratification of salinity and the the Prandtl numberP _r=A_v/K_v, whereA _v andK _v denote the coeffcients of the vertical eddy viscosity and diffusivity, respectively. The two relations result in an extension of the parameter range in which the linear balance of momentum holds.A linear state of motion (LM-state) is defined as the state where the momentum balance is linear. The LM-state comprises the so-called vertically homogeneous and the so-called partially mixed state. Perturbation analysis is introduced and dynamical theory is developed in the LM-state. Since the LM-state covers a fairly wide regime with respect to the balance of salt transport, the state is subdivided into three stages: the diffusive, intermediate and advective stages. In the diffusive stage the upstream salt transport is mainly attributed to the horizontal diffusion, whereas in the advective stage it is attributed to advection caused by gravitational circulation. The salinity balance is also linear in the diffusive and intermediate stages, whereas the balance is nonlinear in the advective stage. The advective stage of the LM-state is regarded as a stage bordering the salt wedge state.The longitudinal distribution of depth-mean salinity is found to take an exponential form in the diffusive stage, a nearly linear form in the advective stage and an intermediate form between them in the intermediate stage.     

Small is beautiful: An inverted view of habitat fragmentation in seagrass beds

An intertidal Zostera marina landscape in Torbay, Devon, UK, was sampled to investigate the relationship between patch size, diversity and infaunal assemblage composition with the intention of defining a minimum Zostera patch size where the infaunal seagrass assemblage becomes distinct from the bare sand assemblage. All Zostera patches were found to support a higher level of biodiversity than the surrounding bare sand. However, the size of the Zostera patch had no impact on the level of diversity; it was just the presence or absence of seagrass that made a difference. The sediment and seagrass variables were not significantly different across the range of Zostera patch sizes, indicating that the environment characteristics were homogeneous within the Zostera patches at the patch scale. Multivariate analysis revealed that assemblage composition did vary between the patch types, although the opportunistic polychaete Capitella capitata was present in all patch types and was the most abundant species overall. The presence of opportunistic species and the homogeneity of the Zostera patch variables may be due to the location of this intertidal seagrass bed, which is relatively exposed compared to the locations of other seagrass beds along the south coast of Devon, resulting in a more dynamic and disturbed environment. Nevertheless, our results demonstrate that even small patches of seagrass comprising a few plants support a higher abundance and diversity of infaunal invertebrates than bare sand, indicating that Zostera patches have conservation value whatever their size.     

Constraining tectonic compression processes by reservoir pressure evolution: Overpressure generation and evolution in the Kelasu Thrust Belt of Kuqa Foreland Basin,NW China

The Kuqa Foreland Basin (KFB) immediately south of the South Tianshan Mountains is a major hydrocarbon producing basin in west China. The Kelasu Thrust Belt in the basin is the most favorable zone for hydrocarbon accumulations. Widespread overpressures are present in both the Cretaceous and Paleogene reservoirs with pressure coefficients up to 2.1. The tectonic compression process in KFB resulted from the South Tianshan Mountains uplift is examined from the viewpoint of the overpressure generation and evolution in the Kelasu Thrust Belt. The overpressure evolution in the reservoir sandstones were reconstructed through fluid inclusion analysis combined with PVT and basin modeling. Overpressures at present day in the mudstone units in the Kelasu Thrust Belt and reservoir sandstones of the Dabei Gas Field and the Keshen zone are believed to have been generated by horizontal tectonic compression. Both disequilibrium compaction and horizontal tectonic compression are thought to contribute to the overpressure development at present day in the reservoir of the Kela-2 Gas Field with the reservoir sandstones showing anomalously high primary porosities and low densities from wireline log and core data. The overpressure evolution for the Cretaceous reservoir sandstone in the Kelasu Thrust Belt evolved through four stages: a normal hydrostatic pressure (>12–5 Ma), a rapidly increasing overpressure (∼5–3 Ma), an overpressure release (∼3–1.64 Ma) and overpressure preservation (∼1.64–0 Ma). Overpressure developed in the second stage (∼5–3 Ma) was generated by disequilibrium compaction as tectonic compression due to the uplift of the Tianshan Mountains acted at the northern monocline of KFB from 5 Ma to 3 Ma, which provided abundant sediments for the KFB and caused the anomalously high sedimentation rate during the N₂k deposition. From 3 Ma to 1.64 Ma, the action of tectonic compression extended from the northern monocline to the Kelasu Thrust Belt and returned to the northern monocline of KFB from 1.64 Ma to present day. Therefore, the horizontal tectonic compression was the dominant overpressure mechanism for the overpressure generation in the third stage (∼3–1.64 Ma) and overpressure caused by disequilibrium compaction from 5 Ma to 3 Ma was only preserved in the Kela-2 Gas Field until present day.     

Characterization of vertical mixing at a tidal-front on Georges Bank

Studies of mixing were done at the northern flank of Georges Bank in the summer and autumn of 1988. Two time-series of the evolution and intensity of microstructure were examined over a tidal period in the context of tidal forcing and the evolution of the density and velocity field at the site. From the CTD, ADCP and microstructure observations (EPSONDE) on Georges Bank, several interesting features of the mixing processes were found. High dissipation and diffusivity regions appear near the bottom of the Bank. Turbulence near the bottom is highest in intensity and reaches farthest from the bottom at peak tidal flow and diminishes in intensity and vertical extent as the flow decreases. The thickness of the bottom turbulent layer has its maximum value when the flow is strongest and the stratification is weakest. Characterization of the dissipation rate and turbulent diffusivities in respect to buoyancy frequency N, current shear S, Richardson Number R_i and ε/νN² was done. Dissipation and χ_T showed little dependence on shear or N² but decreased at larger R_i. χ_t was found to be higher in regions of higher N² and increased as ε/νN² increased. K_T, K and K_ν, were all highest near the bottom in excess of 10⁻²m²s⁻¹ and decreased towards the surface. There was little suggestion of a dependence of mixing efficiency on S², R_i or ε/νN², but some indication that Γ decreases with decreasing N².     

Gassy sediment occurrence and properties: Northern Gulf of Mexico

Free gas is ubiquitous at shallow sediment depths of the northern margin of the Gulf of Mexico. Gassy sediment patches are between 250 and 500 m in horizontal size. Often the gassy layers are within 100 m from the sea floor and are only a few meters thick. Both biogenic and thermogenic gas hydrates have been recovered. Stability values of temperature and pressure indicate that hydrates can exist in water depths less than 500 m. Gassy sediment geoacoustic parameter values are not well constrained because of a lack of concurrent measurements of acoustic properties and sediment gas content. For Gulf of Mexico gassy sediment, some reportedin situ values of sound speed are reduced by an order of magnitude below values for water saturated sediments. More commonly, sound speed is reduced from water saturated sediment values by only 15 to 50 percent.     

Biomass Production in PioneerSpartina anglicaPatches: Evidence for the Importance of Seston Particle Deposition

Salt marsh formation can be initiated by the colonization of bare tidal flats by pioneer halophytes such asSpartina anglica. In the present study, growth ofS. anglicain the edge and centre zones of pioneer patches colonizing a tidal flat in a marine bay in the south-west Netherlands was investigated. Average biomass and shoot length was significantly lower in centre zones than in edge zones of the patches. Multiple linear regression showed that 84% of the variance in aboveground biomass in the centre zones could be explained by sediment phosphorus and nitrogen concentrations. No such relationship existed in the edge zone. This suggests that biomass production in the centre zones of the patches is nutrient limited, but that factors other than sediment nitrogen and phosphorus content determine growth ofSpartinain the edge zones.The sediment nitrogen and phosphorus concentrations in the centre zones of the patches were significantly related with sediment carbon concentrations, indicating that the pool of particle-bound nutrients in the sediment to an important extent is associated with organic particles. Determination of stable carbon isotope signatures of the sedimentary organic matter showed that an increasing carbon content of the sediment in the centre zone of the patches (and a rise in plant biomass) coincides with a higher contribution of non-Spartinaderived carbon to the sedimentary organic matter pool. Sustained biomass production in the centre zones of the patches, thus, probably depends on deposition of allochthonous organic particles and the nutrient inputs inherent in this process. Due to the construction of large-scale hydro-engineering works, however, conditions for deposition of waterborne particles are currently unfavourable. The consequences of reduced carbon and nutrient inputs into the patches will be particularly in the (older) centre zones, where the nutrient pools in non-refractory organic matter have been drained in the years of previous growth. Accordingly, it is hypothesized that the canopy-enhanced sedimentation for whichSpartinais well- known, via positive feed-back effects on plant growth can be crucial for plant vitality, and, hence, for successful colonization of unvegetated areas.     

南海北部2011年台风“纳沙”尾迹处近惯性内波特征的研究

In September 2011, Typhoon Nesat passed over a moored array of instruments recording current and temperature in the northern South China Sea（SCS）. A wake of baroclinic near-inertial waves（NIWs） commenced after Nesat passed the array. The associated near-inertial currents are surface-intensified and clockwise-polarized. The vertical range of NIWs reached 300 m, where the vertical range is defined as the maximum depth of the horizontal near-inertial velocity 5 cm/s. The current oscillations have a frequency of 0.709 9 cycles per day（cpd）, which is 0.025 f higher than the local inertial frequency. The NIWs have an e-folding time-scale of 10 d based on the evolution of the near-inertial kinetic energy. The depth-leading phase of near-inertial currents indicates downward group velocity and energy flux. The estimated vertical phase velocity and group velocity are 0.27 and 0.08 cm/s respectively, corresponding to a vertical wavelength of 329 m. A spectral analysis reveals that NIWs act as a crucial process to redistribute the energy injected by Typhoon Nesat. A normal mode and an empirical orthogonal function analysis indicate that the second mode has a dominant variance contribution of 81%, and the corresponding horizontal phase velocity and wavelength are 3.50 m/s and 420 km respectively. The remarkable large horizontal phase velocity is relevant to the rotation of the earth, and a quantitative analysis suggests that the phase velocity of the NIWs with a blue-shift of 0.025 f overwhelms that of internal gravity waves by a factor of 4.6.     

In vivo metabolism of aromatic amines by the bay mussel, Mytilus edulis

The hazard posed to aquatic organisms by exposure to aromatic amines will depend on their ability to activate or detoxify these potentially genotoxic compounds. We studied the metabolism of four aromatic amines (p-toluidine, aniline, 2-acetylaminofluorene (2-AAF) and 2-aminofluorene (2-AF)) in Bay mussels (Mytilus edulis) to gain an understanding of their metabolic capabilities in vivo. Mussels rapidly accumulated these compounds from the water column and depurated the majority of their steady-state body burdens as metabolites and unmetabolized parent compounds. The biotransformation pathways of [¹⁴C]-labeled parent compounds were determined by analyses of depurated metabolites and tissue residues. Biotransformation of p-toluidine and aniline resulted in the formation of their corresponding N-acetylated detoxification products. Mussels converted small amounts of 2-AAF into a deacetylated product (2-AF) and a mutagenically active metabolite (N-hydroxy-2-AF). 2-AF was extensively detoxified via N-acetylation, which limited the amount of primary amine available for activation.     

Analysis of primary production in the Seto Inland Sea, Japan, using a simple ecosystem model

A simple model of lower trophic level ecosystem has been created to analyze possible environmental control of primary production in eight sub-areas of the Seto Inland Sea. The primary production rates observed by Hashimotoet al. (1997a) in these sub-areas are well reproduced by the model, including horizontal processes such as horizontal transport of nutrients and vertical processes such as vertical mixing, light intensity and sinking of particulate matter. Without taking account of horizontal processes the model also successfully reproduces the observed primary production rates in some areas, but if fails to reproduce those in the others. This shows that the relative importance of the horizontal transport on the primary production differs are by area. Two time scales,T _z andT _H, are introduced to explain this difference.T _z is a vertical cycling time of material, which is defined as the time during which the stock of the material in the water column is utilized for primary production;T _H is the horizontal transit time of the material. The relative importance of the horizontal process is well explained by theT _H/T _z ratio; that is, the horizontal transport process is important in the areas where this ratio is small. Further the possible mechanisms of nutrient supply for the primary production in each sub-area are investigated using this model.