A model for measuring leaf growth rate of vegetative shoot in the seagrass, Zostera marina

A new model for determining leaf growth in vegetative shoots of the seagrass Zostera marina (eelgrass) is described. This model requires the weights of individual mature and immature whole leaves and leaf plastochrone interval (P_L) as parameters, differing from the conventional leaf marking technique (CLM) that requires cutting and separation between new and old tissue of leaves. The techniques required for the model are the same as for the plastochrone method, but the parameters differ between both methods in use of the weight of individual immature leaves. In a mesocosm study, eelgrass growth was examined, and parameters for the new model and plastochrone method (the weights of individual mature and immature leaves and P_L) were measured. Leaf growth rate was measured using the CLM and determined by the new method and the plastochrone method. The results were then compared between the CLM, the new model, and the plastochrone method. The results obtained with the new model were similar to those obtained with the CLM. However, the results of the plastochrone method differed from those of the CLM, while the weight of immature leaves varied seasonally. The new model was also used to determine leaf growth in a natural eelgrass bed in Mikawa Bay, Japan, and revealed the growth rates in all shoots and those of different ages. This method would be advantageous as an accurate means of direct measurement in fieldwork, and should therefore be a useful tool for monitoring seagrass growth.     

A discrete model for estimating the development time from egg to infecting larva of Ostertagia ostertagi parametrized using a fuzzy rule-based system

Ostertagia ostertagi is a nematode, predominantly affecting cattle in the Pampean region of Argentina. A mathematical model parametrized using fuzzy rule-based systems of the Takagi-Sugeno-Kant type (FTSK) for estimating the development time from egg to infecting larval stage L3 of the gastrointestinal parasite O. ostertagi is here proposed. The estimation of development time of O. ostertagi is essential for the generation of appropriate control mechanisms, since this provides information about the time when parasites are ready to migrate to pastures. For the purpose of reflecting the natural environmental conditions, the mean daily temperature is taken as the main and only regulator of the development time. Humidity conditions are considered to be sufficient for the normal development of the larvae. Hence the individual's daily growth is a function of its length and the mean temperature recorded on the previous day. It is expressed in terms of a difference equation with fuzzy parameters, which are defined using laboratory data. Model outputs are tested against results of field experiments. Simulation results are very satisfactory, yielding a mean estimation error (MEE) of 0.64 weeks, with variance 0.34, and a determination coefficient R² = 0.74. The model clearly exhibits an inverse relationship between development time and temperature both in controlled and in field conditions. It also exhibits a very sensitive response both to the order in which the temperature sequence occurs, - reproducing the differences observed between spring and autumn - and to the amplitude of the temperature range.     

An individual-based model to study the reproduction of egg bearing copepods: Application to Eurytemora affinis (Copepoda Calanoida) from the Seine estuary,France

Limited empirical studies have elucidated the daily egg production and associated reproductive processes of egg bearing copepod. Herein, we present an individual-based model which constitutes a realistic representation of the reproduction in egg bearing copepods. The model has been parameterized using an extensive set of experimental data obtained from the literature and from the laboratory and field experiments on the estuarine copepod Eurytemora affinis. The proposed model takes into account the adult female longevity, the clutch size and interclutch duration, which is a function of egg maturation time and latency time required by the female after egg hatching to produce a new clutch. The embryonic development time and hatching success are also taken into account. The effect of temperature on the means and variances of above-mentioned reproductive parameters has been also incorporated. A multi agent system based generic platform “Mobidyc” has been used to generate and calibrate the model. The model demonstrates the reproductive parameters of females of E. affinis which is validated through individual based experiments. Temperature specific simulations provide a dynamical explanation of temperature effect on the cumulative egg production. The daily survival principally affects the number of clutches produced per female during its life span. The results obtained in the present study by combining temperature and survival effects reveal the relatively greater importance of the first factor on the daily egg production of egg-carrying copepods. The present model is generic and hence easily applicable to other animals with comparable reproductive strategy.     

Modeling the population dynamics of cotton bollworm Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) over a wide area in northern China

The cotton bollworm Helicoverpa armigera (Lepidoptera: Noctuidae) is one of the most serious crop pests in northern China, calling for accurate prediction of pest outbreaks and strategies for pest control. A computer model is developed to simulate the population dynamics of H. armigera over a wide area in northern China. The area considered covers 12 provinces where serious outbreaks of H. armigera have been observed. In this model, pest development is driven by local ambient temperature, and adults migrate long distances between regions and select preferred hosts for oviposition within a region. Six types of host including cotton, wheat, corn, peanut, soybean and a single category composed of all other minor hosts are considered in this model. Survival rates of eggs and larvae are based on life-table data, and simulated as a function of host type, host phenology and temperature. The incidence of diapause depends on temperature and photoperiod experienced during the larval stage. Survival rate of non-diapause pupae is a nonlinear function of rainfall, and overwinter survival rate is a nonlinear function of temperature. Insecticide is applied when population density exceeds the economic threshold on a host crop within a region. Comparisons of model output with light-trap data indicate that our model reflects the pest population dynamics over a wide area, and could potentially be used for testing novel pest control strategies in northern China.     

Simulating regeneration and vegetation dynamics in Mediterranean coniferous forests

This study aims to provide a quantitative framework to model the dynamics of Mediterranean coniferous forests by integrating existing ecological data within a generic mathematical simulator. We developed an individual-based vegetation dynamics model, constrained on long-term field regeneration data, analyses of tree-rings and seed germination experiments. The simulator implements an asymmetric competition algorithm which is based on the location and size of each individual. Growth is parameterized through the analysis of tree-rings from more than thirty individuals of each of the three species of interest. A super-individual approach is implemented to simulate regeneration dynamics, constrained with available regeneration data across time-since-disturbance and light-availability gradients. The study concerns an insular population of an endemic to Greece Mediterranean fir (Abies cephalonica Loudon) on the island of Cephalonia (Ionian Sea) and two interacting populations of a Mediterranean pine (Pinus brutia Ten.) and a more temperate-oriented pine (Pinus nigra Arn. ssp. pallasiana) on the island of Lesbos (NE Aegean Sea), Greece. The model was validated against plot-level observations in terms of species standing biomass and regeneration vigour and adequately captured regeneration patterns and overall vegetation dynamics in both study sites. The potential effects of changing climatic patterns on the regeneration dynamics of the three species of interest were subsequently explored. With the assumption that a warmer future would probably cause changes in the duration of cold days, we tested how this change would affect the overall dynamics of the study sites, by focusing on the process of cold stratification upon seed germination. Following scenarios of a warmer future and under the current model parameterization, changes in the overall regeneration vigour controlled by a reduction in the amount of cold days, did not alter the overall dynamics in all plant populations studied. No changes were identified in the relative dominance of the interacting pine populations on Lesbos, while the observed reduction in the amount of emerging seedlings of A. cephalonica on Cephalonia did not affect biomass yield at later stages of stand development.     

Bioenergetics of ephyra larvae of the scyphozoan jellyfish Aurelia aurita in relation to temperature and salinity

The body mass of Aurelia aurita ephyrae was better correlated with the diameter of the central disc than with the distance between opposite rhopaliae or distance between opposite lappet tips. Body dry weight (y, in μg) related to the disc diameter (x, in mm) through the equation y = 22.33 x ^1.99. The exponent 1.99 was significantly lower than that for the medusa stage, indicating a tendency to grow in diameter rather than in weight through the ephyra stage. The average ash-free dry weight (AFDW) of ephyrae was 38.0% of the dry weight. The AFDW/diameter relationship was used to convert measured diameters to body AFDW and calculate succession in body mass, daily ration, daily growth rate and gross growth efficiency. Effects of temperature (6, 9.5, 12, 15 and 18 °C) and salinity (17.5, 22, 26, 30.5 and 35 PSU) on these parameters and feeding were studied at saturated prey concentration (222 Artemia nauplii l⁻¹ initial concentration) by daily measurements over 10 d. There was a strong effect of temperature for total ingestion, growth rate, growth efficiency and final body mass of individual ephyrae, whereas the daily ration was not significantly different between the different temperatures. The experimental group kept at the highest temperature (18 °C) diverged the most, and ephyrae at this temperature ingested 2.7 times more and increased in weight 5.4 times more than at 6 °C. The average daily growth rate and gross growth efficiency of these ephyrae were 34.5% and 25.1%, respectively, significantly higher than at 6, 9.5 and 15 °C. Significant effects of salinity were shown for total ingestion, daily ration, daily growth rate and final weight, although only total ingestion and daily ration diverged sufficiently to show effects in a post-hoc test. This test showed that total ingestion was significantly different for all salinities except between 22 and 35 PSU and between 17.5 and 26 PSU. The daily ration for 35 PSU diverged from all other salinities, whereas none of the other salinities showed any significant differences. Thus, provided food in excess A. aurita can double its weight every 2 to 4 d, dependent on temperature and can therefore develop to the medusa stage in short time. Differences in environmental salinity in the range 17.5 to 35 PSU have little or no effect on growth rate and growth efficiency, whereas our results indicate that the full seawater salinity (35 PSU) causes significantly higher ingestion rate compared to lower salinities. Received: 11 January 1999 / Accepted: 11 May 1999     

Influence of low and decreasing food levels on Daphnia-algal interactions: Numerical experiments with a new dynamic energy budget model

Based on numerical experiments with a new physiologically structured population model we demonstrate that predator physiology under low food and under starving conditions can have substantial implications for population dynamics in predator-prey interactions. We focused on Daphnia-algae interactions as model system and developed a new dynamic energy budget (DEB) model for individual daphnids. This model integrates the κ-rule approach common to net assimilation models into a net-production model, but uses a fixed allocation of net-productive energy in juveniles. The new DEB-model agrees well with the results of life history experiments with Daphnia. Compared to a pure κ-rule model the new allocation scheme leads to significant earlier maturation at low food levels and thus is in better agreement with the data. Incorporation of the new DEB-model into a physiologically structured population model using a box-car elevator technique revealed that the dynamics of Daphnia-algae interactions are highly sensitive to the assumptions on the energy allocation of juveniles under low food conditions. Additionally we show that also other energy allocation rules of our DEB-model concerning decreasing food levels and starving conditions at the individual level have strong implications for Daphnia-algae interactions at the population level. With increasing carrying capacity of algae a stable equilibrium with coexistence of Daphnia occurs and algae shifts to limit cycles. The amplitudes of the limit cycles increase with increasing percentage of sustainable weight loss. If a κ-rule energy allocation is applied to juveniles, the stable equilibrium occurs for a much narrower range of algal carrying capacities, the algal concentration at equilibrium is about 2 times larger, and the range of algae carrying capacities at which daphnids become extinct extends to higher carrying capacities than in the new DEB-model. Because predator-prey dynamics are very sensitive to predator physiology under low food and starving conditions, empirical constraints of predator physiology under these conditions are essential when comparing model results with observations in laboratory experiments or in the field.     

A multi-agent simulation to assess the risk of malaria re-emergence in southern France

A multi-agent simulation (MAS) was developed to assess the risk of malaria re-emergence in the Camargue in southern France, a non-endemic area where mosquitoes of the genus Anopheles (Culicidae) live. The contact rate between people and potential malaria vectors, or the human biting rate, is one of the key factor to predict the risk of re-emergence of malaria, would the parasite be introduced in the region. Our model (called MALCAM) represents the different agents that could influence malaria transmission in the Camargue – people, mosquitoes, animal hosts and the landscape – in a spatially explicit environment. The model simulates spatial and temporal variations in human biting rate at the landscape scale. These variations depend on the distribution of people and potential vectors, their behaviour and their interactions. A land use/cover map was used as a cellular-spatial support for the movements of and interactions between mobile agents. The model was tested for its sensitivity to variations in parameter values, and for the agreement between field observations and model predictions. The MALCAM model provides a tool to better understand the interactions between the multiple agents of the disease transmission system, and the land use and land cover factors that control the spatial heterogeneity in these interactions. It allows testing hypotheses and scenarios related to disease dynamics by varying the value of exogenous biological, geographical, or human factors. This application of agent-based modelling to a human vector-borne disease can be adapted to different diseases and regions.     

Computer simulation applied to the biological control of the insect Aphis gossypii for the parasitoid Lysiphlebus testaceipes

In integrated pest management (IPM), biological control is one of the possible options for the prevention or remediation of an unacceptable pest activity or damage. The success of forecast models in IPM depends, among other factors, on the knowledge of temperature effect over pests and its natural enemies. In this work, we simulated the effects of parasitism of Lysiphlebus testaceipes (Cresson, 1880) (Hymenoptera: Aphidiidae) on Aphis gossypii (Glover, 1877) (Hemiptera: Aphididae), a pest that is associated to crops of great economic importance in several parts of the world. We made use of experimental data relative to the host and its parasitoid at different temperatures. Age structure was incorporated into the dynamics through the Penna model. The results obtained showed that simulation, as a forecast model, can be a useful tool for biological control programs.     

Aphid population response to agricultural landscape change: A spatially explicit,individual-based model

A spatially explicit individual-based simulation model has been developed to represent aphid population dynamics in agricultural landscapes. The application of the model to Rhopalosiphum padi (L.) population dynamics is detailed, including an outline of the construction of the model, its parameterisation and validation. Over time, the aphids interact with the landscape and with one another. The landscape is modified by varying a simple pesticide regime, and the multi-scale spatial and temporal implications for a population of aphids is analysed. The results show that a spatial modelling approach that considers the effects on the individual of landscape properties and factors such as wind speed and wind direction provides novel insight into aphid population dynamics both spatially and temporally. This forms the basis for the development of further simulation models that can be used to analyse how changes in landscape structure impact upon important species distributions and population dynamics.     

Modeling the role of macroalgae in a shallow sub-estuary of Narragansett Bay, RI (USA)

Proliferation of macroalgal mats is a frequent consequence of nutrient-driven eutrophication in shallow, photic coastal marine ecosystems. These macroalgae have the potential to significantly modify water quality, plankton productivity, nutrient cycling, and dissolved oxygen dynamics. We developed a model for Ulva lactuca and Gracilaria tikvahiae in Greenwich Bay, RI (USA), a shallow sub-estuary of Narragansett Bay, as part of a larger estuarine ecosystem model. The model predicts the biomass of both species in units of carbon, nitrogen, and phosphorus as a function of primary production, respiration, grazing, decay, and physical exchange, with particular attention to the effects of biomass layering on light attenuation and suppression of metabolic rates. The model successfully reproduced the magnitude and seasonal cycle of area-weighted and peak biomass in Greenwich Bay along with tissue C:N ratios, and highlighted the importance of grazing and inclusion of self-limitation primarily in the form of self-shading to overcome an order of magnitude difference in rates of production and respiration. Inclusion of luxury nutrient uptake demonstrated the importance of internal nutrient storage in fueling production when nutrients are limiting. Macroalgae were predicted to contribute a small fraction of total system primary production and their removal had little effect on predicted water quality. Despite a lack of data for calibration and a fair amount of sensitivity to individual parameter values, which highlights the need for further autecological studies to constrain formulations, the model successfully predicted macroalgal biomass dynamics and their role in ecosystem functioning. Our formulations should be exportable to other temperate systems where macroalgae occur in abundance.     

Development and validation of an individual based Daphnia magna population model: The influence of crowding on population dynamics

An individual-based model was developed to predict the population dynamics of Daphnia magna at laboratory conditions from individual life-history traits observed in experiments with different feeding conditions. Within the model, each daphnid passes its individual life cycle including feeding on algae, aging, growing, developing and – when maturity is reached – reproducing. The modelled life cycle is driven by the amount of ingested algae and the density of the Daphnia population. At low algae densities the population dynamics is mainly driven by food supply, when the densities of algae are high, the limiting factor is “crowding” (a density-dependent mechanism due to chemical substances released by the organisms or physical contact, but independent of food competition).     

Allowing macroalgae growth forms to emerge: Use of an agent-based model to understand the growth and spread of macroalgae in Florida coral reefs,with emphasis on Halimeda tuna

The growth patterns of macroalgae in three-dimensional space can provide important information regarding the environments in which they live, and insights into changes that may occur when those environments change due to anthropogenic and/or natural causes. To decipher these patterns and their attendant mechanisms and influencing factors, a spatially explicit model has been developed. The model SPREAD (SPatially-explicit Reef Algae Dynamics), which incorporates the key morphogenetic characteristics of clonality and morphological plasticity, is used to investigate the influences of light, temperature, nutrients and disturbance on the growth and spatial occupancy of dominant macroalgae in the Florida Reef Tract. The model species, Halimeda and Dictyota spp., are modular organisms, with an “individual” being made up of repeating structures. These species can also propagate asexually through clonal fragmentation. These traits lead to potentially indefinite growth and plastic morphology that can respond to environmental conditions in various ways. The growth of an individual is modeled as the iteration of discrete macroalgal modules whose dynamics are affected by the light, temperature, and nutrient regimes. Fragmentation is included as a source of asexual reproduction and/or mortality. Model outputs are the same metrics that are obtained in the field, thus allowing for easy comparison. The performance of SPREAD was tested through sensitivity analysis and comparison with independent field data from four study sites in the Florida Reef Tract. Halimeda tuna was selected for initial model comparisons because the relatively untangled growth form permits detailed characterization in the field. Differences in the growth patterns of H. tuna were observed among these reefs. SPREAD was able to closely reproduce these variations, and indicate the potential importance of light and nutrient variations in producing these patterns.     

Effects of food-level acclimation on digestive enzyme activities and feeding behavior of Calanus pacificus

A recent hypothesis in the zooplankton literature states that zooplankton acclimate to ambient food concentrations such that higher digestive enzyme activities and, consequently, higher maximum ingestion rates are achieved at higher food levels. To test this hypothesis, adult female Calanus pacificus, collected from the main basin of Puget Sound, Washington, USA, in August 1979 and May 1982, were conditioned for 2 wk at different concentrations of the diatom Thalassiosira weissflogii (=fluviatilis). Ingestion rates and the activity of the digestive enzymes laminarinase, maltase, and cellobiase were measured periodically during acclimation and in a block-designed feeding experiment at the end of acclimation. Consistent with the hypothesis, maximum ingestion rate and digestive enzyme activity were positively correlated. However, in contrast to the hypothesized mechanism, this result arose because both maximum ingestion rate and digestive enzyme activity were negatively correlated with food concentration during acclimation. The enhanced ingestion of copepods following long-term (12 to 14 d) acclimation to low food is similar to that previously described for short-term (e.g. 1 d) starvation. It might be energetically optimal for copepods experiencing a patchy food environment to maintain higher levels of digestive enzymes at low food concentrations in order to exploit high concentrations of food when encountered.     

Simulating population variation and movement within fragmented landscapes: An application to the gopher tortoise (Gopherus polyphemus)

As the human activity footprint grows, land-use decisions play an increasing role in determining the future of plant and animal species. Studies have shown that urban and agricultural development cannot only harm species populations directly through habitat destruction, but also by destroying the corridors that connect habitat patches and populations within a metapopulation. Without these pathways, populations can encounter inbreeding depression and degeneration, which can increase death rates and lower rates of reproduction. This article describes the development and application of the FRAGGLE model, a spatial system dynamics model designed to calculate connectivity indices among populations. FRAGGLE can help planners and managers identify the relative contribution of populations associated with habitat patches to future populations in those patches, taking into account the importance of interstitial land to migration success. The model is applied to the gopher tortoise (Gopherus polyphemus), a threatened species whose southeastern U.S. distribution has diminished significantly within its native range due to agricultural and urban development over the last several decades. This model is parameterized with life history and movement traits of the gopher tortoise in order to simulate population demographics and spatial distribution within an area in west-central Georgia that supports a significant tortoise population. The implications of this simulation modeling effort are demonstrated using simple landscape representations and a hypothetical on land-use management scenario. Our findings show that development resulting in even limited habitat losses (10%) may lead to significant increases in fragmentation as measured by a loss in the rate of dispersions (31%) among area subpopulations.     

Elevated testosterone advances onset of migratory restlessness in a nearctic-neotropical landbird

The primary cue for initiation of spring migratory restlessness (Zugunruhe) in landbird migrants is photoperiod. Gonadal hormones are known to have a role in the regulation of migratory disposition; however, the extent of their effect is not well understood. We examined the impact of exogenous testosterone on the onset of migratory restlessness in gray catbirds (Dumetella carolinensis). Catbirds were stratified by sex and randomly assigned to two rooms; individuals in one room were photoadvanced to initiate migratory restlessness and the other room was maintained on a 12:12 light/dark photoperiod. Each room had three groups (n?=?10/group); males with testosterone implants, males with empty implants, and females. We predicted that in the photoadvanced room males with testosterone implants would initiate migratory activity earlier than empty-implanted males. We found that in the photoadvanced group, testosterone-implanted males initiated migration 2 weeks prior to empty-implanted males, and 3 weeks prior to females. In the non-photoadvanced males, the testosterone-implanted males initiated migration at the same time as the corresponding group in the photoadvanced room, while the empty-implanted males and females did not exhibit Zugunruhe. Our results illustrate that elevated testosterone can advance the onset of Zugunruhe, even in the absence of an extended photoperiod. Additionally, the onset of migratory restlessness observed in the photoadvanced, non-testosterone-implanted males and females further supports the importance of photoperiod as a cue for migratory restlessness. An interesting observation was the intersexual differences in the onset of migratory activity in gray catbirds, a species not previously known to exhibit protandry.     

Differential effect of ultraviolet exposure (UVR) in the stress response of the Dinophycea Gymnodinium sp. and the Chlorophyta Dunaliella tertiolecta: mortality versus survival

Dunaliella tertiolecta (Chlorophyta) and Gymnodinium sp. (Dinophyceae) cells were exposed to ultraviolet radiation (UVR) (PAR, UVA and UVB: PAB) for 6 days either continuously or during a photoperiod. Both UVR treatments were harmful to Gymnodinium but exposure to continuous PAB had the most dramatic effects. Although a number of lesions/damage could have happened during the first few hours of exposure to UVR, in less than 24 h, Gymnodinium lost its ability to detoxify ROS efficiently, photoinhibition occurred, thymine dimers formed in the DNA, caspase-like enzymatic activities DEVDase sharply increased and cells died as determined by SYTOX-green staining. Superoxide dismutase activity did not significantly change with time, and although the catalase activity augmented in both treatments, cells still suffered from the UVR stress. Clearly, UVR was fatal to the dinoflagellate. For the chlorophyte, however, cell numbers increased regardless of the UVR treatment and mortality remained low (<20 %). F _v/F _m showed an initial decrease but then remained constant for both light treatments. After 6 days of continuous PAB exposure, however, signs of stress (thymine dimers, oxidative stress) paralleled a drop in catalase activity. Results obtained here demonstrate that the dinoflagellate Gymnodinium was much more sensitive and was harmed more rapidly by UVR exposure than the chlorophyte D. tertiolecta. The increased tolerance to UVR exposure of the chlorophyte may provide advantages over other more sensitive phytoplankton species within the photic zone. We provide strong support in the present study for repair being an important component of UV resistance in this species.     

Mathematical analysis of change in forest carbon use efficiency with stand development: A case study on Abies veitchii Lindl.

Changes in carbon use efficiency (CUE), which is defined as the ratio of net primary production (NPP) to gross primary production (GPP), were analyzed for Abies veitchii Lindl. forests with respect to stand development by developing a simple mathematical model incorporating data on physiological variables and leaf mass ratio. A decrease in CUE with stand development was successfully expressed as a function of stand biomass (y) based on the following three assumptions: (1) a power-law relationship between mean respiration and mean individual tree mass, (2) a power-functional relationship between mean gross primary production and mean individual tree mass, and (3) self-thinning relationship between stand biomass and density. Based on this model, a parameter of CUE–y relationship was defined, and it was clarified that CUE decrease with stand development is caused not by the ratio of specific respiration rate to specific gross photosynthetic rate, but by leaf mass ratio. Since CUE is high in young forests, helpful information on selecting woody species when planting seedlings was provided from the viewpoints of reducing CO₂ in the atmosphere and global warming.     

Integrating soil carbon cycling with that of nitrogen and phosphorus in the watershed model SWAT: Theory and model testing

In this paper we describe and test a sub-model that integrates the cycling of carbon (C), nitrogen (N) and phosphorus (P) in the Soil Water Assessment Tool (SWAT) watershed model. The core of the sub-model is a multi-layer, one-pool soil organic carbon (S_C) algorithm, in which the decomposition rate of S_C and input rate to S_C (through decomposition and humification of residues) depend on the current size of S_C. The organic N and P fluxes are coupled to that of C and depend on the available mineral N and P, and the C:N and N:P ratios of the decomposing pools. Tillage explicitly affects the soil organic matter turnover rate through tool-specific coefficients. Unlike most models, the turnover of soil organic matter does not follow first order kinetics. Each soil layer has a specific maximum capacity to accumulate C or C saturation (S_x) that depends on texture and controls the turnover rate. It is shown in an analytical solution that S_x is a parameter with major influence in the model C dynamics. Testing with a 65-yr data set from the dryland wheat growing region in Oregon shows that the model adequately simulates the S_C dynamics in the topsoil (top 0.3 m) for three different treatments. Three key model parameters, the optimal decomposition and humification rates and a factor controlling the effect of soil moisture and temperature on the decomposition rate, showed low uncertainty as determined by generalized likelihood uncertainty estimation. Nonetheless, the parameter set that provided accurate simulations in the topsoil tended to overestimate S_C in the subsoil, suggesting that a mechanism that expresses at depth might not be represented in the current sub-model structure. The explicit integration of C, N, and P fluxes allows for a more cohesive simulation of nutrient cycling in the SWAT model. The sub-model has to be tested in forestland and rangeland in addition to agricultural land, and in diverse soils with extreme properties such high or low pH, an organic horizon, or volcanic soils.