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The development of the DNDC plant growth sub-model and the application
of DNDC in agriculture: A review
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Review
A R T I C L E I N F O A B S T R A C T
Article history:
Received 10 January 2016 Plant growth plays an important role in regulating soil C and N as well as water regimes and can therefore
Received in revised form 8 June 2016 influence soil biochemical or geochemical processes. A sub-model was built in DNDC (DeNitrification-
Accepted 9 June 2016 DeComposition) to simulate crop growth; since its development, it has often been modified and adapted
Available online xxx to suit specific purposes, crops and circumstances. Here, we review the chronological history of various
versions of the DNDC plant growth sub-models and present the results of a literature search regarding
Keywords: the application of the DNDC model to various crops in agriculture. We found that food, oil and sugar crops
Process model were the primary research focus and accounted for 67.5%, 12.5% and 6.3% of all DNDC crop-based studies,
DNDC
respectively. We also summarize the research achievements published in recent years, and conclude that
Crop types
the DNDC plant growth sub-model could be successfully used to assist in predicting trace gas emissions
Agriculture
Greenhouse gas and soil carbon and nitrogen dynamics after modifying some of the parameters obtained from relevant
literature to suit local cultivars. The objective of this study is to provide DNDC users with an
understanding of the model mechanisms related to field and horticultural crops, with suggestions for
modelling different crops and outlining further model applications and modifications.
ã 2016 Elsevier B.V. All rights reserved.
Contents
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271
2. Plant growth sub-model evolution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 272
3. The role of plant growth sub-models . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 274
4. Application of DNDC for various crop types . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275
4.1. Model validation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275
4.2. Food crops . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 276
4.3. Oil crops . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 276
4.4. Sugar crops . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 276
4.5. Other crops . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277
5. Discussion: DNDC model use and development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277
5.1. Strengths and weaknesses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277
5.2. Prospects for model development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
6. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
1. Introduction
http://dx.doi.org/10.1016/j.agee.2016.06.017
0167-8809/ã 2016 Elsevier B.V. All rights reserved.
272 Y. Zhang, H. Niu / Agriculture, Ecosystems and Environment 230 (2016) 271–282
and exhibit high degrees of temporal and spatial variability (Smith used internationally (Li et al., 2003, 2005b; Babu et al., 2005; Tang
et al., 2008a; Giltrap et al., 2010). Agriculture releases substantial et al., 2006; Wang et al., 2008).
amounts of CO2, CH4 and N2O into the atmosphere (Cole et al., Fig. 1 Within agro-ecosystems, plant growth plays a crucial role
1997; IPCC, 2001; Paustian et al., 2004). Carbon dioxide is primarily in regulating soil C and N and water regimes and can therefore
released through microbial decay or by the burning of plant litter influence the biochemical or geochemical processes (see Fig. 1).
and soil organic matter (Janzen, 2004; Smith, 2004); methane is Plant growth is also a crucial component of any ecosystem model
produced when organic materials decompose in oxygen-deprived as plants remove water, N and other nutrients from soils and
conditions, for example, during fermentative digestion by rumi- consequently can alter a series of biogeochemical reactions. Plants
nant livestock or in stored manures and rice grown under flooded additionally produce biomass, including yield and litter, which
conditions (Mosier et al., 1998); and nitrous oxide is generated by provide food security and improve soil resources. Nevertheless,
the microbial transformation of nitrogen in soils and manures. N2O developed crop sub-models are not included in most existing
production is often enhanced when available N exceeds crop ecosystem models, and most models developed by agronomists are
requirements, especially in wet conditions (Smith and Conen, highly parameterized, which is useful for field-scale yield studies
2004; Oenema et al., 2005). Agricultural greenhouse gas (GHG) but not widely applicable at a regional scale (Grant et al., 1993;
fluxes are complex and heterogeneous, but the active management Grant, 1995; Parton et al., 1998; Liu et al., 2000; Li, 2002 Grant,
of agricultural systems offers possibilities for mitigation. 1995; Parton et al., 1998; Liu et al., 2000; Li, 2002). To address this
As GHG emissions are highly sensitive to many factors, gap, a plant growth sub-model was built in DNDC to simulate the
including soil type, climate conditions and management practices, effects of cumulative temperature, N uptake, and water stress on
high degree of uncertainty is associated with GHG emissions. crop growth at a daily time step (Li et al., 1994). The basic rules in
Direct measurement of greenhouse gas emissions for inventory this sub-model of the original DNDC model were not newly
purposes is impractical as it would require a prohibitively large invented but rather adopted from existing plant growth models.
number of measurements over large areas for long periods of time. This sub-model forms a bridge between plant growth and C and N
Thus, the development of a more process-based approach is biogeochemical cycles.
desirable. To date, process-based models have been used to Currently, improvements to the DNDC plant growth sub-model
estimate agricultural GHG-mitigation potential by comparing are neither well documented nor widely understood either by the
alternative agricultural management scenarios at local and research community or potential users. To rectify this and to
national scales (Williams et al., 1992; Lokupitiya and Paustian, integrate existing data with prior knowledge, this review
2006; Desjardins et al., 2010; Shepherd et al., 2011; Cui et al., 2014). summarizes the state of the DNDC plant growth sub-model. In
DNDC (DeNitrification-DeComposition) is a process-based particular, this review (1) explores and describes the main
model of carbon and nitrogen biogeochemistry in agro-ecosystems developments of different DNDC versions and their plant growth
that was originally developed to simulate N2O emissions from sub-models, (2) reviews the huge variety of alternative
cropped soils in the United States (Li et al., 1992a). From the time of approaches, (3) assesses model application in a variety of crop
its initial development, classical laws of physics, chemistry and types and (4) highlights the strengths, weaknesses and potential
biology, as well as empirical equations generated from laboratory future improvements of the model. Additionally, this review
studies, have been incorporated in this model to parameterize each provides a theoretical basis for a comprehensive evaluation of the
specific geochemical or biochemical process, and numerous eco-environmental effects of various crops using a DNDC-based
changes have been made to the DNDC model to develop country- model.
or need-specific models (Li et al., 1992b; Frolking et al., 1999; Salas
et al., 2005; Li, 2007; Giltrap et al., 2010). The DNDC model can now 2. Plant growth sub-model evolution
be used to simulate crop growth, soil temperature and moisture
regimes, soil carbon dynamics, nitrogen leaching, and the In the early stages of model development, DNDC versions did
emissions of greenhouse and trace gases and has been widely not contain a plant growth sub-model (Li et al., 1992a). Until 1994,
Fig. 1. Generation and consumption of nitrous oxide (N2O), carbon dioxide (CO2) and methane (CH4) in forest/arable soils and the relationships among these three fluxes
(modified from Fang et al., 2014).
Y. Zhang, H. Niu / Agriculture, Ecosystems and Environment 230 (2016) 271–282 273
the DNDC model (Version 7.1) was supplemented with an direct and diffuse light separately (Spitters, 1986; Spitters et al.,
empirical plant growth sub-model that contained sub-routines 1986). The integration of photosynthesis rate with time and canopy
for land cropping practice and management to study the profile is conducted using the three-point Gaussian integration
biogeochemistry of soil C in arable land; some major crops could method (Goudriaan, 1986). The response of photosynthesis to light
be simulated in this model (Watts and Hanks, 1978; Li et al., 1994). is expressed as an exponential function with two parameters
In this version of the DNDC plant growth sub-model, (1) nitrogen (Penning de Vries et al., 1989). The effects of temperature on
uptake by vegetation is the key process linking crop growth with photosynthesis are simulated as influencing photosynthesis rate
climate and soil status. The daily N uptake rate is regulated by four through light saturation and initial light use efficiency (Penning de
factors: the crop potential maximum yield, which is defined as the Vries et al., 1989). The effect of atmospheric CO2 concentration on
optimum grain yield of a crop growing with sufficient water and photosynthesis rate is simulated based on Goudriaan et al. (1984);
nitrogen and could be calculated using formulas detailed in Li et al. photosynthesis is also influenced by water and nitrogen stress
(1994); the crop C/N ratio, which is the default in Li et al. (1994); factors. (4) Respiration and assimilate allocation: Crop respiration
the generalized crop growth curve and a similar curve only for is simulated considering growth and maintenance respiration
winter wheat, which are used for all crops (Watts and Hanks, separately (McCree, 1979; Penning de Vries et al., 1989). The
1978); and the availability of dissolved inorganic nitrogen (NO3 difference between gross photosynthesis and respiration is the
and NH4+) in the soil profile. The proportion of N uptake from the amount of assimilate available for allocation among crop organs.
NO3 and NH4+ pools is determined by their relative abundances; Assimilate allocation is simulated based on phenological stages
(2) crop biomass carbon is calculated at harvest as the product of (Jones and Kiniry, 1986; Ritchie et al., 1988; Penning de Vries et al.,
actual annual N uptake and the fixed crop C/N ratio. The total 1989). If the assimilate is not sufficient for the growth requirement
biomass carbon is allocated to grain, straw/stalks, and roots; and of the grain, the deficiency will be translocated from the stem;
(3) root respiration is a result of three processes: root growth, root otherwise the remainder will be allocated to the stem (Yoshida,
maintenance, and ion uptake and transport (Veen, 1981). 1972; Gao et al., 1992). (5) Rooting process: Rooting processes
Later, many changes made to produce the PnET-N-DNDC model include the increase of root front depth, the distribution of root
were incorporated into DNDC (Li et al., 2000; Li, 2000). In this plant length density and biomass in soil profile. In the model, the
growth sub-model of DNDC (Version 8.0), by integrating crop deepening rate of the root front is proportional to the thermal time
characteristics, climate, soil properties, and farming practices, the before flowering, and root front depth is limited to a maximum
plant growth sub-model simulated plant growth and its effects on depth (1 m). Daily variation in root length density in a layer
soil systems driven by solar radiation, temperature, water stress, depends on new root growth and root senescence. Root biomass
and N stress and passed litter production and water and N distribution in the soil profile is estimated based on root length
demands to the soil climate or decomposition sub-models. In this distribution, which follows an exponential pattern in the soil
sub-model, some farmland management measures, such as profile, but it is subjected to the influence of constraint factors for
fertilization, have been parameterized. By application of ammoni- each layer, including a static factor and four dynamic factors
um, nitrate, or anhydrous ammonia to the soil, the nitrogen is (Ritchie et al., 1988; Jones et al., 1991). (6) Water and nitrogen
directly allocated into the corresponding inorganic N pools, uptake: Crop water uptake depends on potential transpiration
whereas in the case of urea application, the model will first demand determined by LAI and climate conditions and on uptake
simulate the hydrolysis of urea (Butterbach-Bahl et al., 2001). capacity determined by soil moisture, root length and root
The DNDC model was further modified by integrating several distribution in soil. Water stress factor is estimated based on
key crop algorithms, which were developed as a component of the ratio of actual water uptake and potential transpiration
Crop-DNDC, to produce a phenological crop growth sub-model demand (Ritchie et al., 1988). Crop nitrogen uptake depends on
that could simulate crop growth by tracking physiological crop demand and uptake capacity. Nitrogen uptake capacity
processes, i.e., calculating water stress and nitrogen stress (Li, depends on mineral nitrogen concentration in the root zone and
2000; Zhang et al., 2002a). Phenological stages and stress factors soil moisture, which are simulated by soil biogeochemical and
(water and nitrogen) influence carbon allocation and nitrogen hydrological components. Nitrogen demand and uptake capacity
demand. The major processes of the crop sub-model include are simulated based on formulas (Godwin and Jones, 1991).
phenological development, leaf area index (LAI), photosynthesis More recent versions of DNDC have incorporated processes
and respiration, assimilate allocation, rooting processes, and water from other models, such as Photosynthesis-Evapotranspiration
and nitrogen uptake: (1) Phenological development: the life cycle (PnET), Economic Farm Emission Model (EFEM) and Northern
of crops is divided into nine phenological development stages Ecosystem Soil Temperature (NEST), or have revised some
based on CERES models (Ritchie et al., 1987, 1988; Ritchie, 1991). parameters to assume the basic structure of a complex modular-
Active crop growth stages range from emergence to maturity, and based model, such as Forest-DNDC, EFEM-DNDC and NEST-DNDC.
phenological development rate is simulated based on thermal time A number of models have also been developed for specific regions
(Jones and Kiniry, 1986; Ritchie, 1991), and the thermal time (e.g., NZ-DNDC, DNDC-Europe) or specific crops (e.g., DNDC-Rice,
needed from sowing to emergence is estimated based on sowing DNDC-CSW) (Rahn et al., 2012).
depth. For other stages, the required thermal time is an input To explicitly describe photosynthesis, respiration, and C
genetic parameter or estimated based on the thermal time of the allocation in crops, a generic crop model, MACROS, was modified
former stages, whether for wheat or for rice and maize (Ritchie and integrated with DNDC v. 9.0 (Penning de Vries et al., 1989). In
et al., 1987; Ritchie, 1991; Kiniry, 1991). (2) Leaf area index (LAI): this integration, net carbon gain was calculated by subtracting the
LAI variation is simulated as the difference between leaf area respiration requirement from canopy photosynthesis. This assimi-
growth (associated with assimilate allocation) and leaf senescence lated carbon was allocated among plant organs using partitioning
(associated with phenological development and stress factors) for coefficients, which were defined as functions of the developmental
wheat or for rice and maize (Jones and Kiniry, 1986; Ritchie et al., stage (e.g., transplanting stage, heading stage, or maturation
1988; Kropff and van Laar, 1993). LAI growth is then simulated stage). Carbon flux from plant roots to the soil in the form of
according to the assimilate allocation. Leaf senescence is estimated respiration, turnover of organic matter, which was assumed to be
based on phenological stages and the effects of water and nitrogen 1% of root biomass that was lost daily to the soil after the heading
stress factors (Jones and Kiniry, 1986; Ritchie et al., 1988). (3) stage, and exudation, which occurred at all phenological stages,
Photosynthesis: Gross photosynthesis is simulated considering were all components of the carbon balance for a given crop
274 Y. Zhang, H. Niu / Agriculture, Ecosystems and Environment 230 (2016) 271–282
(Fumoto et al., 2008). New crop types or cultivars (e.g., deep-water vegetables, papaya, potato, beet, paddy rice, and banana (Smith
rice) can be added to the model by defining their physiological and et al., 2010; Deng et al., 2011a; Li et al., 2012a).
phenological parameters. With these mechanisms, DNDC is able to Over the years, a regionalized Canadian version of DNDC (DNDC
predict the impacts of alternative crop rotations or new cultivars v. CAN1), which is modified from the DNDC v. 9.5 release version,
on GHG emissions from the wetland crop fields. With the above has been utilized to better represent crop production, soil C and
described modifications, DNDC has become capable of simulating GHG emissions for cool weather cultivars of maize, soybeans and
the fundamental biogeochemical processes occurring in agro- wheat in Canada (Kröbel et al., 2011; Smith et al., 2013 Smith et al.,
ecosystems across climate zones, soil types, and management 2013).
regimes. The chronological development of plant growth sub-models in
In BE-DNDC, a special tool, “Crop Creator”, was developed and DNDC versions is detailed in Fig. 2 and Appendix A.
integrated into the later versions of DNDC. “Crop Creator” is a crop
parameter calculator that allows users to permanently modify the 3. The role of plant growth sub-models
parameter values stored in the crop library of DNDC for existing
crops based on their own observations. “Crop Creator” also allows The current input interface in DNDC for crop data allows the
users to create new crop types and calculate the biomass, C pools, definition of (1) types of crops consecutively or simultaneously
C/N ratio, total N demand and other parameters of the new crop planted in a given year, (2) planting and harvest dates, (3) crop
that are routinely reported by farmers or agronomists. Physiologi- residue management, and (4) crop physiological and phenological
cal or phenological factors can be more closely approximated to parameters (see Table 1). All of the default values provided by
observed values (Beheydt et al., 2007). DNDC can be modified to adjust the actual crop heat/water/N
Recent versions of DNDC have also included simulations of crop demands, growth curves, biomass partitioning or yield. After
growth and alternative farming management practices, such as the defining this group of crop parameters, the crop growth sub-model
use of nitrification and urease inhibitors, slow-release fertilizers, forms the first component of DNDC, in conjunction with the soil
sprinkler and drip irrigation, and plastic film mulching, so that they climate and decomposition sub-models, to predict soil tempera-
may be used in GHG mitigation studies. DNDC provides a wide ture, moisture, pH, redox potential (Eh) and substrate concentra-
spectrum of data to help assess the best management practices tion profiles driven by ecological drivers (e.g., climate, soil,
based on their impacts on a relatively complete span of ecosystem vegetation and anthropogenic activity). Then, this sub-model
services. The latest version of DNDC is version 9.5, in which some of combines with the second component of DNDC, consisting of the
the past developments regarding crop modelling are no longer nitrification, denitrification and fermentation sub-models, to
usable and detailed phenological model is also not included predict emissions of carbon dioxide (CO2), methane (CH4),
(Gilhespy et al., 2014). For instance, the phenological growth model ammonia (NH3), nitric oxide (NO), nitrous oxide (N2O) and
for crops in DNDC are still in the code but are commented out, and dinitrogen (N2) from the plant-soil systems. The modelled daily
a simpler GDD-based model is employed; the leaf area index photosynthesis, respiration, C allocation, and water and N uptakes
becomes an estimated model output and is not involved in crop are recorded so that the modelled results can be checked against
growth simulation in the model. A total of 63 crop types can now be observations to ensure that the crops are simulated correctly. Crop
simulated in cropping systems in the model, including fallow, demand for N is calculated based on the optimum daily crop
maize, winter wheat, soybean, legume hay, non-legume hay, growth and the plant C/N ratio. The actual N uptake by a crop could
spring wheat, sugarcane, barley, oats, alfalfa, sorghum, cotton, rye, be limited by N or water availability during the growing season.
Fig. 2. Schematic diagram of the DeNitrification-DeComposition (DNDC) and the major improvements of its plant growth sub-model (modified from Li et al., 1994, 2006; Li,
2000; Butterbach-Bahl et al., 2001; Zhang et al., 2002a).
Y. Zhang, H. Niu / Agriculture, Ecosystems and Environment 230 (2016) 271–282 275
Table 1
Main inputs to DNDC for crop physiological and phenological parameters (modified from the latest DNDC manual).
Parameter Description
Crop type Select one of the crop types parameterized in DNDC or select “New_crop” from the list to add new crops.
Fraction of leaves and stems left in the field The fraction of the above-ground crop residue left in the field after harvest.
after harvest
Maximum biomass, kg C ha 1 The maximum biomass production for grain, leaves, stems and roots under optimum growing conditions. The default is
that 1 kg dry matter contains 0.4 kg C.
Biomass fraction The grain, leaf, stem and root fractions of the total biomass at maturity.
Biomass C/N ratio The ratio of C/N for grain, leaves, stems and roots.
Total N demand, kg N ha 1 Total amount of N required by the crop to reach maximum production.
Thermal degree days, C Cumulative air temperature from seeding until maturity of the crop.
Water demand, g water/g dry matter Amount of water required for the crop to produce one unit of biomass dry mass.
N fixation index The ratio of total N content in the plant and plant N taken from the soil for legume crops; the default is 1 for non-legume
crops.
LAI adjustment factor A factor for adjusting default specific leaf weight.
Optimum temperature, C The optimum crop growth temperature.
This N is extracted from the available soil NO3 and NH4+ pools (in the plant growth sub-models. In addition, other information was
proportion to the relative size of each pool) to the depth of the collected from these articles, including record keeping of previous
plant roots. DNDC partitions the crop biomass production model versions and code changes, validation of previous model
increment among the leaves, stems, roots and grain based on versions, collaborations and publications, and the intended use of
the plant growth stage at each daily time step. After harvest, all the model, e.g., an assessment of the impact of land use change or
root biomass remains in the soil profile, and a fraction of the above- an economic analysis.
ground crop residue also remains on top of or in the soil profile
until next tilling application. The crop residue incorporated into 4.1. Model validation
the soil will be partitioned into three soil litter pools, namely very
labile, labile and resistant litter pools, based on its C/N ratio. The Model validation against experimental data is essential to verify
litter incorporation provides essential input for the soil organic that the model is correctly simulating the underlying processes. To
matter (SOM) storage and hence integrates the plant and soil into a date, DNDC has been used to simulate various crop types and
biogeochemical system. Thus, by taking up water and N from and cropping systems in many countries for many purposes. Giltrap
depositing litter into the soil, plant growth regulates the soil water, et al. (2010) has reviewed some model validation studies of DNDC
C and N regimes, which in turn determine a series of biochemical and found that agreement between the model simulations and
reactions, including C sequestration and trace gas emissions. More measured values vary. Therefore, this review only introduces
details are found in the DNDC manual. model validation studies that used the latest version of DNDC (see
Table 2).
4. Application of DNDC for various crop types Overall, DNDC has proven effective in characterizing the general
influences of different conditions and farmland management
A literature search was conducted for DNDC-based models practices on GHG emissions, soil moisture and temperature, plant
published since 1992 and cited by SCI and SCIE. There are 80 growth and crop yields, and N leaching losses. In addition, the
articles that describe in total more than 20 types of crops in their simulation of evapotranspiration has become a research focus
sample plots, covering varieties of rotation methods. This literature since the Penman-Monteith equation has been integrated with
review was used to document the chronological development of DNDC to calculate reference evapotranspiration (Dutta et al., 2016).
Table 2
Validation studies comparing predictions against measurements using the latest version of DNDC. “” means overestimation; “ ” means underestimation.
4.2. Food crops that the simulated results of DNDC are helpful in choosing best
management practices to decrease nitrate leaching and N2O and
Fig. 3 shows that studies that utilize DNDC-based models NO emissions (Ludwig et al., 2011; Smith et al., 2013; Cui et al.,
primarily focus on the following crops: rice, wheat, maize, barley, 2014; Sansoulet et al., 2014). In addition, DNDC is also widely used
potatoes and other food crops; studies of these crops account for and compared with other models to investigate the influence of
67.5% of all DNDC crop-based studies. Three primary reasons exist future climate change and the application of advanced agricultural
for the prevalence of food crops in DNDC crop-based studies. (1) management measures to achieve good application results in
Alimentary crops are the main source of human food, and their barley fields (Abdalla et al., 2011; Smith et al., 2012; Cardenas et al.,
cultivation covers approximately 85% of the cropping area, 2013; Balashov et al., 2014; Deng et al., 2015).
globally. Simulation of greenhouse gas emissions from soils used
for food crops is highly relevant to global climate change. (2) The 4.3. Oil crops
DNDC model was developed in agricultural nations, and historical
in-situ observations of food crops are also abundant in these areas; Studies focusing on oil crops (i.e., primarily rapeseed and
this correspondence provides a theoretical basis for the validation soybean) account for 12.5% of all DNDC crop-based studies. Oil
and further application of the DNDC model. (3) DNDC has been crops are mainly used in the production of food, feed, machinery,
widely verified in food crop research, and some crop-specific metallurgy, soap, paint, rubber, plastics and pharmaceuticals, and
models (e.g., DNDC-CSW and DNDC-Rice) have also been devel- they play an important role in the global economy. Thus, a great
oped (Grant et al., 2004; Liu et al., 2006; Tonitto et al., 2007a,b; deal of scientific research has been conducted on the effect of
Zhang et al., 2009; Li et al., 2012b). climate change on oil crops (Wardlaw et al., 1980). In studies
Rice, maize, wheat and barley are the main food crops that are conducted in soybean fields, Lugato et al. (2010) used the DNDC
frequently simulated by DNDC. Recent studies conducted in rice model with the objective of improving the national GHG inventory
paddy fields under different soil conditions and agricultural in Italy to build a platform of simulation for the entire national
management measures suggest that DNDC and DNDC-Rice can territory; Ouyang et al. (2013) employed DNDC to predict the long-
be used to estimate GHG emissions, aboveground biomass and term dynamics of nitrous oxide (N2O), carbon dioxide (CO2) and
total global warming potential (GWP) (Zhang et al., 2011; methane (CH4) under different crop patterns and cultivation
Katayanagi et al., 2012, 2013; Fumoto et al., 2013). In maize fields, management practices in an agricultural freeze zone in China and
the related DNDC studies are mainly focused on N cycling and finally confirmed that the manure amendment and N fertilizer
conclude that the farmland management measures significantly application can improve the SOC, increase crop production and
affect the nitrogen fluxes and nitrate leaching amounts (Follador enhance the GWP.
et al., 2011; Perlman et al., 2013; Jarecki et al., 2015; Chen et al.,
2015; Zhang et al., 2015). Several studies have investigated the 4.4. Sugar crops
effectiveness of the DNDC model to access and predict GHG
emissions and crop yields under different management practices Sugar crops hold an important position in global agriculture and
and emissions scenarios in wheat fields and have demonstrated the global economy. Studies focusing on sugar crops account for
Fig. 3. Frequency of DeNitrification-DeComposition (DNDC) model use for different crop types. Crop types in rotations were not included in this figure. The type designated as
“other” includes various vegetables and fruits.
Y. Zhang, H. Niu / Agriculture, Ecosystems and Environment 230 (2016) 271–282 277
6.3% of all DNDC crop-based studies. Sugar beets and sugarcane defined, which increases model options (Gilhespy et al., 2014). (3)
crops are the main sugar crops in the world. The sugar beet is a The professional simulation functions allow the simulation of crop
temperate crop, and its cultivation area is mainly distributed in the growth, climate, soil organic matter decomposition, nitrification,
Northern Hemisphere. Europe is the largest beet producing area, denitrification, and fermentation, and DNDC describes carbon and
accounting for more than 75% of global sugar beet production. nitrogen circulation and provide comprehensive and detailed
Sugarcane is mainly cultivated in tropical and subtropical regions; output data reported at a daily time step for most parameters
Brazil, India and China are the primary sugarcane-producing (Zhang et al., 2002a,b). (4) DNDC accurately simulates N2O
countries in the world. Sugar crop studies that utilize DNDC emissions at the mechanistic level and calculate greenhouse gas
models mostly focus on the sugar beet due to the wide application emissions from agro-ecosystems (Li et al., 2004b, 2007). (5) A
of DNDC models in temperate regions (Lugato et al., 2010). Abdalla sophisticated regional simulation can be performed using GIS data.
et al. (2013) conducted a study that supported the use of the DNDC DNDC can simulate greenhouse gas emissions at the provincial or
model to predict the consequences of climate change on Net national scale with the support of a GIS database (Li et al., 2005b;
Ecosystem Exchange (NEE) from sugar beet fields in south-eastern Qiu et al., 2011; Chirinda et al., 2011 Chirinda et al., 2011). (6) DNDC
Ireland and suggested that the soil carbon decreased at the arable also provides tools to quantify the uncertainties for site or regional
site with the contribution of exported carbon; this arable simulations. With the Monte Carlo approach at the site or grid cell
ecosystem was predicted to operate as a carbon sink in simulated scale, and with the MSF (most sensitive factor) method at regional
climate change scenarios through 2060. scale, the uncertainties produced from the model simulations can
be quantified (Deng et al., 2011a).
4.5. Other crops Although DNDC is frequently used to calculate GHG emissions,
they have several limitations. (1) DNDC is a 1-D model that does
Currently, the relevant studies conducted in other economic not simulate lateral flows of water and nutrients and assumes a
crop fields focus on GHG emissions. Kwack and Kobayashi (2011) homogenous microbial population for both autotrophs and
applied the DNDC model for estimating the impacts of farm heterotrophs (Smith et al., 2008a,b), thus the DNDC model may
management practices on climate change and soil properties in the simulate well the overall magnitude of N2O emissions but not
tea fields in Japan. The use of compost significantly reduced nitrate simulate well the timing of individual events (Saggar et al., 2004;
leaching and N2O emissions in comparison with N fertilizer, and Smith et al., 2008b). (2) In cold locations, DNDC inaccurately
changes in soil pH, soil texture, temperature and precipitation simulates annual soil N2O emissions where snow cover, freeze and
affected N2O emissions, nitrate leaching, and SOC sequestration. Yu thaw may create anaerobic conditions and cause microbes to die of
and Zhao (2015) used the DNDC model to simulate the temporal frost (Butterbach-Bahl et al., 2001; Smith et al., 2004; de Bruijn
variation in soil respiration in a cotton field in China and found a et al., 2009; Kim et al., 2012), suggesting that the mechanism of ice-
good correlation between predicted and measured root respiration trapped N2O release in the DNDC model is not quite correct; this
and an underestimation of the cumulative CO2 emissions. Zhang has been partially addressed by model modifications and evolution
et al. (2016) proved the DNDC model to be a powerful tool for (Kariyapperuma et al., 2011). (3) The model does not consider the
addressing the efficacy of alternative management practices in effect of soil depth on N2O diffusion rate and only simulates a
vineyards in Ningxia, China and revealed that N2O emissions can be heterogeneous soil profile to 50 cm in depth, which may cause the
reduced by adopting drip irrigation systems rather than traditional model to calculate slightly earlier times for N2O emissions peaks
furrow irrigation systems. than are observed in the field (Li et al., 1992a,b). However, with the
assumption of a deep water pool located below the bottom of the
5. Discussion: DNDC model use and development modelled soil profile (50 cm) and above the drainage tiles (145 cm)
to control the tile discharge flow, this soil depth (0–50 cm) should
Crop simulation plays a crucial role in modelling C and N be deep enough to cover the most important biogeochemical
biogeochemistry in agro-ecosystems. With the addition of the processes while minimizing computing time (Li et al., 2006). (4)
plant growth sub-model, the DNDC model is well adapted to DNDC will underestimate soil moisture if observed soil moisture
simulate harvested yield and biomass as well as N uptake and GHG remains above field capacity because DNDC uses a simple cascade
emissions under different agricultural management practices and (tipping bucket) water flow model that drains the profile to field
land-uses. Meanwhile, the strengths and weaknesses of the DNDC capacity (Kröbel et al., 2010; Uzoma et al., 2015). Likewise, because
plant growth sub-model are well illustrated by the application of DNDC does not include unsaturated flow and underestimates the
the DNDC model to farmlands, as discussed below. rainfall loss caused by surface runoff and leaf interception, it may
overestimate soil moisture at other times (Kiese et al., 2005;
5.1. Strengths and weaknesses Tonitto et al., 2010). Improvements to the soil hydrology
framework of the model should be considered for further studies.
Compared with most C & N models, DNDC is built using more (5) The predicted N2O emissions from organic manure in the
detailed processes, and their utility comes from having a modelling absence of any additional N fertilization are often too low (Abdalla
framework that simulates a large array of interacting nutrient et al., 2009, 2011). To fix this error, a portion of the N in manure is
flows and emissions as well as ease of use and moderate input allocated to various N pools in Manure-DNDC and the other most
requirements. DNDC primarily possesses several advantages. (1) recent DNDC models, with a sizable amount going directly to the
DNDC is written in C++, and thus its input and output interfaces are NH3 pool. The recent models still require further testing with
convenient to operate. (2) A wide variety of available inputs exist, respect to manure applications.
as does a comprehensive library of default settings, which enables Although some deficiencies remain, DNDC offers users consid-
users to properly parameterize the crop cultivars and soils and to erable flexibility, not only in the types of situations modelled but
model a wide range of sites and situations without the need for also in the output available for evaluation (Li et al., 2001; Sleutel
large quantities of rarely measured input data (Grosz et al., 2015). et al., 2006). The DNDC model is currently applied to various
Input parameter data (soil, climate, and agricultural activity) are ecosystems worldwide. The willingness of DNDC model devel-
easy to acquire, and a geographic information system (GIS) opers to collaborate with users to improve the model and its
database can also be obtained by converting national statistical performance is a notable strength that is deserving of recognition.
data. Furthermore, many input parameters can also be user-
278 Y. Zhang, H. Niu / Agriculture, Ecosystems and Environment 230 (2016) 271–282
The strengths of the DNDC model suggests that it will continue to dimensional model to conduct spatial and temporal simulation
be readily applied in the future (Li et al., 2014). of biomass accumulation, soil carbon and nitrogen dynamics, and
GHG emissions of various crops in the future.
5.2. Prospects for model development
Acknowledgments
As a process-based model, DNDC can both model the
environmental impacts of agricultural management systems and The authors are indebted to Prof. Changsheng Li at the Earth
improve our understanding of the underlying processes (Ange- Systems Research Center, University of New Hampshire, Durham,
nendt, 2003; Babu et al., 2006; Dou et al., 2014). In regional mode, NH 03824, for providing valuable suggestions. DNDC would not be
DNDC can be used to develop national inventories, assess the an excellent process-based model without the years of concen-
changes in greenhouse gas emissions in relation to the expected trated efforts of the original DNDC developer, Prof. Li, and other
changes in management and climate, and verify mitigation researchers. The authors would also like to thank Yao Li at the
strategies (Frolking et al., 2002; Li et al., 2002; Cai et al., 2003; College of Resources and Environment, University of Chinese
Hastings et al., 2010; Wang et al., 2011). Academy of Sciences, Beijing, and Jing Ge at the Shanghai Institutes
By incorporating comments and suggestions from users for Biological Sciences, Chinese Academy of Sciences, Shanghai, for
worldwide, the DNDC model suite has continued to be improved assisting with the data processing and visualization. This work was
over the several decades of its development. For example, one supported by the Asian Development Bank project (RETA6422).
recent study integrated two fundamental hydrologic features (the
Soil Conservation Service curve function and the Modified Appendix A.
Universal Soil Loss Equation) with DNDC to simulate the
movement of water and nutrients in the horizontal and vertical
dimensions at the watershed or landscape scales and demonstrat- Chronological development of the versions of the plant growth sub-
ed that DNDC could contribute more effectively to watershed N model of DeNitrification-DeComposition (DNDC). This table, which is
loading studies by incorporating basic hydrological features into modified from a table published in a review, complements the text
the model framework (Deng et al., 2011a,b). This innovation descriptions of the model versions with an at-a-glance chronological
allowed the modified DNDC to quantify N loading at the watershed summary of the main characteristics and developments within the
scale. Further exploration of watershed carbon and nitrogen cycles plant growth sub-models of different versions of DNDC (Gilhespy et al.,
and the joint management practices of animal husbandry and 2014).
cultivation are expected to become vital components of DNDC
research in the future. Publication Model Model description and plant growth sub-
Some large scale model testing, development and intercom- version model evolution
parison projects are ongoing, such as AGMIP (Agriculture Model Li et al. (1992a) DNDC v. Consisted of three sub-models: (1) soil-
Intercomparison and Improvement Project), GRA (Global Research 1.0–7.0 climate/thermal-hydraulic flux, (2)
Alliance) and its modelling platform GRAMP, and FACCE-JPI (Food decomposition (three SOC pools), (3)
denitrification (nitrification only present
Security and Climate Change). With the objective of improving the as a simple equation). No crop growth
predictions of soil carbon and nitrogen cycling in the context of sub-model.
climate change, these projects provide places where researchers Li et al. (1994) DNDC v. 7.1 Modified the decomposition sub-model
can share information regarding biogeochemical ecosystem (four SOC pools) and added an empirical
plant growth sub-model to calculate daily
modelling and connect with other researchers. This sharing of
root respiration, N uptake by plants, and
research and ideas can improve the DNDC model for use as a plant growth. The major crops that could
benchmark in the simulations of crop growth and C & N dynamics. be simulated in the model were maize,
soybean, winter wheat, spring wheat,
6. Conclusions barley, sorghum, alfalfa, non-legume hay,
cotton, and sugarcane.
Li et al. (2000) PnET-N- A process based model that integrated
DNDC plant growth sub-models have undergone many changes, DNDC three existing models, namely, the
and the continued importance of DNDC to scientific research is Photosynthesis-Evapotranspiration
reflected by its current number of users, its vast literature base, and (PnET) model, DNDC and the nitrification
model, adapted for temperate forest
the large number of model versions. Through a review of various
ecosystems.
versions of the DNDC plant growth sub-models and the research Li (2000); Li et al. DNDC v. 8.0 Added a new two-component model
products regarding the application of DNDC models to various (2000) framework (nitrification and
crops in agriculture, we concluded that the DNDC plant growth fermentation), an ‘anaerobic balloon’, and
sub-models could effectively simulate the biogeochemical process the effect of freezing and thawing (as
developed in PnET-N-DNDC). By
in crop growth, although DNDC is not a crop model, and found that
integrating crop characteristics, climate,
food, oil and sugar crops were the primary research focus for 67.5%, soil properties, and farming practices, the
12.5% and 6.3% of all DNDC crop-based studies, respectively. DNDC plant growth sub-model simulated plant
has been widely applied for these field crops, which mainly growth and its effects on soil system.
Zhang et al. (2002a) Crop-DNDC Integrated detailed crop growth
included rice, wheat, maize, barley, rapeseed, soybean and sugar
algorithms that coupled soil
beets, with minor modifications of model parameters to suit local biogeochemical components with the
cultivars, and achieves reasonable simulation results. However, for DNDC model. Crop growth was simulated
horticulture and other crops, DNDC has not been modelled and not only by tracking crop physiological
continually validated. These related studies still need to be processes but also by calculating water
stress and nitrogen stress. The simulated
completed and challenged.
crop types included wheat, rice and
Some researchers have used DNDC integrated with spatial maize.
models to conduct regional and global simulation. The DNDC Zhang et al. (2002a) DNDC v. 8.2 A new phenological crop sub-model, as
model (and particularly the plant growth sub-model) is being developed in Crop-DNDC, was introduced
continuously improved and may be recreated as a two-
Y. Zhang, H. Niu / Agriculture, Ecosystems and Environment 230 (2016) 271–282 279
(Continued) (Continued)
Publication Model Model description and plant growth sub- Publication Model Model description and plant growth sub-
version model evolution version model evolution
as an alternative to the empirical crop various types of pure forests (young to
growth sub-model. old, and including a variety of species).
Zhang et al. (2002b) Wetland- Integrated the PnET-N-DNDC and the Smith et al. (2010) DNDC v. 9.3 Improvement to estimates of soil
DNDC FLATWOODS models (Sun et al., 1998) for evaporation under different levels of
use in wetland ecosystems. surface residue cover. A total of 62 crop
Brown et al. (2002) UK-DNDC Some alteration of the database structure types were simulated in the model,
and content of DNDC was required to including fallow, maize, winter wheat,
improve the model’s suitability for soybean, legume hay, non-legume hay,
application in the UK. A total of 18 crop spring wheat, sugarcane, barley, oats,
types were simulated, including winter alfalfa, sorghum, cotton, rye, vegetables,
wheat, winter barley, spring barley, oats, potato, beet, paddy rice, banana, and
potatoes, vegetables, sugar beets, and celery.
rapeseed. Kröbel et al. (2011) DNDC-CSW Introduction of an empirical sub-model to
Li et al. (2004a) DNDC v. 8.5 Modification of the ‘anaerobic balloon’ DNDC v. 9.3. The Canadian Spring Wheat
concept to incorporate the Nernst and (CSW) sub-model included a modified
Michaelis-Menten equations (from plant biomass growth curve, dynamic
Wetland-DNDC). plant C/N ratios and modified plant
Li et al. (2004b) DNDC-Rice A generic crop model, MACROS, was biomass fractioning curves to allow a
modified and integrated with DNDC to more accurate estimation of spring wheat
simulate the development and growth of growth and N uptake in Canadian agro-
several crops, including rice (Penning de ecosystems.
Vries et al., 1989). New crop types or Haas et al. (2012) Landscape- The soil biogeochemical processes of the
cultivars (e.g., deep-water rice) and DNDC agricultural DNDC and the Forest-DNDC
simulations of rice crop rotations were were combined in a general soil
now included in this model. Eleven biogeochemistry model to simulate
dominant rice-cropping systems were ecosystem C and N turnover and changes
added to the database. in soil C and N stocks for various land-use
Saggar et al. (2004) NZ-DNDC Modifications were made to DNDC v. 6.7 types and periods of land-use change.
to better represent New Zealand’s grazed Zhang et al. (2012) NEST-DNDC Integration of DNDC with Northern
pasture systems. Ecosystem Soil Temperature (NEST) to
Li et al. (2005a) Forest- Constructed by integrating two existing permit the modelling of the interactions
DNDC models, PnET and DNDC, with several of soil thermal-hydrological conditions
new features for upland and wetland and biogeochemical processes in
forest ecosystems. permafrost soils. NEST-DNDC maintained
Kiese et al. (2005) Forest- PnET-N-DNDC modified for tropical the capacity to model upland and wetland
DNDC- rainforest ecosystems. ecosystems without permafrost.
Tropical Li et al. (2012a) Manure- Modification of DNDC to describe organic
Neufeldt et al. (2006) EFEM- A GIS-coupled economic-ecosystem DNDC matter turnover and gas emissions for
DNDC model incorporating the Economic Farm manure based on manure systems at the
Emission Model (EFEM) (Angenendt, farm scale. Approximately fifty crops (e.g.,
2003) and DNDC v. 8.0. maize, wheat, soybean, alfalfa, sorghum,
Li et al., 2006); DNDC v. 9.0 Improved simulation of free ammonium oats, barley, legume hay, non-legume hay,
Fumoto et al. dynamics, nitrification, and nitrate and cover crops) were parameterized in
(2008) leaching. Manure-DNDC.
Beheydt et al. (2007) BE-DNDC Integration of DNDC v. 8.3 P with regional Li et al. (2012a) DNDC v. 9.4 Shares the same soil NH3 algorithms as
data for Belgium. A special tool, “Crop developed for the Manure-DNDC. Crop
Creator” was developed in this version. types include maize, wheat, soybean,
“Crop Creator” is a crop parameter alfalfa, sorghum, oats, barley, legume hay,
calculator that allows users to non-legume hay, and cover crops.
permanently modify the parameter Li, pers comm. 2013 DNDC v. 9.5 Improvements to crop growth
values stored in the crop library of DNDC simulations, hydrological features and
for existing crops based on their own other improvements to meet the demand
observations. “Crop Creator” also allows for GHG mitigation studies. A total of 63
users to create new crop types and crop types were simulated in the model,
calculate the biomass, C pools, C/N ratio, including fallow, maize, winter wheat,
total N demand and other parameters of soybean, legume hay, non-legume hay,
the new crop that are routinely reported spring wheat, sugarcane, barley, oats,
by farmers or agronomists. Physiological alfalfa, sorghum, cotton, rye, vegetables,
or phenological factors can be more papaya, potato, beet, paddy rice, and
closely approximated to observed values. banana. In this most recent version, crop
Leip et al. (2008) DNDC- Integration of CAPRI (Common types were slightly different than in
Europe Agricultural Policy Regional Impact previous versions.
Assessment) (Britz, 2005) and DNDCv.8.9 Kröbel et al. (2011) DNDC v. Modification of DNDC95 to better
in Europe. A total of 29 crops were CAN1 represent cool weather cultivars of maize,
simulated in this model, and their crop soybeans and wheat in Canada.
parameters were obtained from the Dutta et al. (2016) DNDC v. Integration of DNDC v. CAN1 to correct
DNDC default library (Li et al., 2004b). CAN2 the reference evapotranspiration per crop
Grote et al. (2009); Mobile- PSIM and DNDC were previously linked type by multiplying the reference
Grote et al. (2011) DNDC by plant uptake of nitrogen. The DNDC evapotranspiration, determined through
model that was used in the MoBiLE the previous Penman-Monteith equation,
framework was able to calculate “water by a crop coefficient Kc, based on the crop
pools and fluxes throughout the total type and growth stage.
rooted soil profile with soil layer specific
parameterizations”. MoBiLE-DNDC was
capable of simulating carbon fluxes in
280 Y. Zhang, H. Niu / Agriculture, Ecosystems and Environment 230 (2016) 271–282
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