Agricultural .Sjmw~.~. Vol. 52, Nns 213, pp.

171 198, 1996

CopyrIght t 1996 Published by Elsewer Science Ltd
Printed in Great Britain. All rights reserved
0308-521X:96 $15 00’ 00
ELSEVIER PII: SO308-521X(96)OOOll-X

The ‘School of de Wit’ Crop Growth Simulation Models:

A Pedigree and Historical Overview

B. A. M. Bournan,” H. van Keulen,” H. H. van Laarh

& R. Rabbingeh

“DLO-Research Institute for Agrobiology and Soil Fertility, P.O. Box 14.
Wageningen 6700 AA, The Netherlands

‘Department of Theoretical Production Ecology, Agricultural University, Wageningen.

The Netherlands

(Received 8 March 1996)

ABSTRACT

In this paper, a pedigree of rhe crop growth .simulation modeis by. the
‘School of de Wit’ is presented. The origins and philosophy of this school
we trtlced,from de Wit’s classical publication on modelling photosynthesis
of leqf canopies in 1965. It is shown how changing research gouls und
priorities over the years have resulted in the evolution of u pedigree of‘
models that are similar in philosophy hut d@er in level of complexit!,, the
processes addressed and their,functionalityl. In the beginning, modelling HUT
motivated by the quest for scienttjic insight and the lvi.4 to quantify. and
integrate hiophysical processes to explain the observed vuriution in crop
gro\vth. Later, the emphasis of undfundingjor, agricultural research shifted
to\zlardsputting ucquired insights to practical and operutional use. Model
development became led by a demand,for tacticul und strategic decision sup-
port, ?,iell,fi,recusting, land zonation and explorative scenario studies. Model-
ling developments ,for dtyerent production situutions are illustrated using
the models the uuthors consider most important, i.e. BACROS, SD’C’ROS,
WOFOST, MACROS and LINTEL, but reference is also made to other
models. Finully, comments we made about the usefulness and ~pplicuhilitj’of’
these models qfter nearly 30 years of’tkveiopment. and some,future cour.ses
of action are suggested. Copyright t$ I996 Published h,, Elsrvier Science Lttl

INTRODUCTION

By the end of the 196Os, computers had evolved sufficiently to allow and
even to stimulate the first attempts to synthesize detailed knowledge on
171
172 B. A. M. Bouman, H. van Keulen, H. H. van Laar, R. Rabbinge

plant physiological processes, in order to explain the functioning of crops

as a whole. Insights into various processes were expressed using math-
ematical equations and integrated in so-called simulation models. These
first (heuristic) models were meant to increase the understanding of crop
behaviour by explaining crop growth and development in terms of the
underlying physiological mechanisms. Over the years, new insights and
different research questions motivated the further development of simula-
tion models. In addition to their explanatory function, the applicability of
well-tested models for extrapolation and prediction was quickly recog-
nized, and more application-oriented models were developed. For
instance, demands for advisory systems for farmers and scenario studies
for policy makers resulted in the evolution of models geared towards
tactical and strategic decision support, respectively (Rabbinge, 1986; van
Keulen & Penning de Vries, 1993). Now, crop growth modelling and
simulation have become accepted tools for agricultural research (Rab-
binge, 1986; Seligman, 1990). A wide variety of crop models has been
developed all over the world to serve many different purposes, with major
modelling groups in the USA in the former project IBSNAT (Interna-
tional Benchmark Sites Network for Agrotechnology Transfer) (Uehara &
Tsuji, 1993; Tsuji et al., 1994) in Australia with the system APSIM
(Agricultural Production system SIMulator) (McCown et al., 1995) and
in The Netherlands at Wageningen. In Wageningen, crop growth model-
ling was initiated and developed by the late C. T. de Wit (deceased 1993)
and his co-workers at the Department of Theoretical Production Ecology
of the Wageningen Agricultural University (TPE-WAU) and the DLO-
Research Institute for Agrobiology and Soil Fertility (AB-DLO; and
CAB0 and IBS) (de Wit, 1970). Since then, many scientists have followed
in his footsteps and have taken up crop modelling. In response to chan-
ging research goals and policies over the years, a range of crop models has
emerged that often confuses the outsider who is ‘merely’ looking for the
Wageningen crop growth model. In this paper, therefore, a historical
overview is given of the pedigree and developments of the ‘School of de
Wit’ models, with a brief description of some of the most significant
models. The overview is limited to dynamic simulation models for growth
and development of field crops I. Scientific details of the various models
are not given here as they have been amply described in books and litera-
ture referenced in this paper (e.g. the series of ‘Simulation Monographs’,

‘A more complete compilation of (European) agro-ecosystem models was recently started

within the framework of the concerted action for the development and testing of quanti-
tative methods for research on agricultural systems and the environment (CAMASE)
(Plentinger & Penning de Vries, 1995).
 Crop growth simulation models 113

published by PUDOC between 1971 and 1993). Finally, the authors pre-
sent a personal review of the usefulness and applicability of these models
after more than 30 years of development.

SYSTEM AND MODEL CHARACTERISTICS

A model is a simplified representation of a system, and a system is a lim-

ited part of reality that contains interrelated elements (de Wit, 1982~). The
system we consider here is the agricultural cropping system. In 1982, de
Wit and Penning de Vries proposed a classification of this system into four
production situations:
Production situation I: Potential production. Growth occurs in conditions
with ample supply of water and nutrients and growth rates are determined
solely by weather conditions (solar radiation and temperature).
Production situation 2: Water-limited production. Growth is limited by
shortage of water during at least part of the growing period but nutrients
are in ample supply.
Production situation 3: Nitrogen-limited production. Growth is limited
by shortage of nitrogen (N) during at least part of the growing season, and
by water or weather conditions for the rest of the time.
Production situation 4: Nutrient-limited production. Growth is limited
by a shortage of phosphorus (P), or of other minerals for at least part of
the growing season, and by N, water or weather conditions for the rest.
In all four situations, pests, diseases or weeds may further reduce crop
yield. In practice, actual production situations are difficult to assign to any
of these four situations, but this practical simplification of schematizing
specific situations allows progress to be made, particularly at the start of a
study (de Wit & Penning de Vries, 1982; Rabbinge, 1986; van Duiven-
booden & Gosseye, 1990). Recently, a new classification of agricultural
production systems was introduced in the C. T. de Wit Graduate School of
Production Ecology (Rabbinge, 1993): potential growth is defined by the
concentration of atmospheric COZ, solar radiation, temperature and crop
characteristics; attainable growth is determined by the limiting factors of
water and nutrients; actual growth is reduced below the attainable by fac-
tors such as weeds, pests, diseases and pollutants (Fig. 1). The development
of models for each of these situations proceeded at its own rate and in its
own direction, depending on the research goals and objectives at the time.
Technically, most models of the ‘School of de Wit’ are characterized by
the labels ‘dynamic’, ‘hierarchical’, ‘state-variable based’, ‘explanatory’
and ‘deterministic’. They are dynamic because rates of change in the
system (e.g. growth rate) are calculated as a function of time, using time
174 B. A. M. Bouman, H. van Keulen, H. H. van Laar, R. Rabbinge

I I
/I
VI
1.5 I’ 5 10 ton ha-’

Production level

Fig. 1. The relationship among potential, attainable and actual yield and defining, limiting
and growth reducing factors (Rabbinge, 1993).

coefficients that are typical for the processes that are described (de Wit,
1982b). The time coefficient, which has units of time, is the inverse of the
characteristic rate of a process. This inclusion of time differentiates them
from static models in which, for example, crop production is statistically
regressed on weather variables. Second, they divide the system under
study into hierarchical levels of organization, e.g. cells, organs, plants,
crop. These hierarchical levels exhibit characteristic behaviours that result
from the integration of lower-level processes (Loomis et al., 1979). For
instance, a leaf light-response characteristic is the result of processes at the
lower levels of cells and chloroplasts (Sinclair et al., 1977), and canopy
photosynthesis is the sum of photosynthesis of all the individual leaves (de
Wit, 1965). Different hierarchical levels can be combined in one model
provided that time-coefficients that are appropriate for each level of hier-
archy are used (de Wit, 1970). Mathematical modelling entails quantita-
tive integration of the mechanisms at the various hierarchical levels
to provide an explanation of system behaviour. Third, the system is
characterized by a set of state variables (e.g. weights) that are updated at
each iteration or time-step, by rate variables (e.g. carbon flux). The time-
step is typically one quarter of the time coefficient. Values of the rate
variables are calculated from information about the current state of the
system and from external, environmental (e.g. solar radiation) and
auxiliary variables (e.g. leaf area index) (Fig. 2). Fourth, the models are
explanatory because the calculations involving rate variables are based on
 Crop growth simulation models 175

solar air
irradiation temperature
I I

structural
biomass
stor. organs

roots
I

I rate I
Fig. 2. Diagram of the relations in a typical “School of de Wit” crop growth model
(SUCROS) for potential production. Boxes indicate state variables, valves rate variables.
circles auxiliary variables, solid lines (arrows) the flow of matter and dotted lines the flow
of information.

knowledge of the underlying physical, physiological and biochemical pro-

cesses. Only when knowledge is lacking, or a simplification is required, are
descriptive, i.e. statistical, relationships used. However, within a hier-
archical structure, descriptive relationships at lower levels become expla-
natory at higher levels (Loomis et al., 1979). Fifth, the dynamic simulation
models of the ‘School of de Wit’ are deterministic because all plants in the
crop are considered to be of the same genotype, and exposed to the same
initial and environmental (soil, weather) conditions. Crop characteristics
and environmental conditions are therefore expressed as a single set of
model parameter values and external model input data, respectively. The
176 B. A. M. Bouman, H. van Keulen, H. H. van Laar, R. Rabbinge

TABLE 1
Steps in the Conceptual Phases of Model Development: Conceptual or Preliminary,
Comprehensive and Summary

Conceptual or preliminary phase

1 Formulation of objectives
2 Definition of system limits
3 Conceptualization of the system (states, rates, auxiliary variables, forcing vari-
ables etc.)
Comprehensive phase

4 Quantification through literature, process experiment or estimation of the relation

between rate and forcing variables, states or auxillary variables
5 Model construction (definition of computer algorithm)
6 Model verification, i.e. testing the intended behaviour of the model
Summary phase

7 Evaluation of model performance, i.e. testing the model in parts or as a whole,

using independent experiments on system level
8 Sensitivity analysis (numerical or structural)
9 Simplification
IO Formulation of decision rules or forecasting models to be used for practical
applications

After Rabbinge & de Wit (1989).

apparently stochastic nature of real biological systems, expressed in

genetic, temporal and spatial variation, can be mimicked using numerical
techniques such as Monte Carlo simulation (Klepper & Rouse, 1991;
Bouman, 1994; Rossing et al., 1994).
In describing the developments of the crop growth models of the ‘School
of de Wit’, we follow the classification of development phases introduced
by Penning de Vries (1980) who distinguished preliminary, comprehensive
and summary models. These three phases of model development are
described in Table 1. Preliminary, or conceptual, models reflect current
scientific knowledge, and are simple in structure because of incomplete
knowledge of the component processes. With increasing insight,
preliminary models can evolve into comprehensive models that represent
systems in which the essential elements are thoroughly understood and
which contain large amounts of information. In plant physiological
research the main purpose of both preliminary and comprehensive models
is to formalize and integrate knowledge of plant growth processes, to test
hypotheses by comparing model results with experiments, to structure
research programmes and to extrapolate from the laboratory to the field.
In brief, the aim is to increase our understanding of crop performance.
After the first comprehensive models had been built and tested, however,
 Crop growth simulation models 177

they were quickly recognized as powerful tools for exploring situations and
possibilities of crop production that were almost impossible to investigate
using the conventional methods and techniques of experimentation.
Because comprehensive models are typically complex and hardly accessible
to potential users, this stimulated the development of summary models. A
summary model can be regarded as a model of a (comprehensive) model, in
which essential elements are simplified and aspects that are only marginally
important are ignored. Summary models have typically been developed
in response to application-oriented research questions (e.g. tactical and
strategic). Their use, for example, may be in decision support systems for
pest and disease management and for plant nutrient management.

MODEL RESPONSES TO CHANGING RESEARCH QUESTIONS

The early years (1965-1980): gaining insight

Crop modelling evolved in the late 1960s as a means of integrating

knowledge about plant physiological processes in order to explain the
functioning of crops as a whole. Researchers set themselves the goal of
quantifying qualitative speculations about the effects of canopy structure,
solar radiation and transpiration on canopy photosynthesis. The outcome
of their work was a set of preliminary and comprehensive models for
production situations 1 and 2.

Potential production situation: ELCROS and BACROS

In 1965, de Wit published the classic report ‘Photosynthesis of leaf canopies’
(Fig. 3), in which a procedure was described ‘that allowed calculation of
daily photosynthesis of a canopy with known characteristics for any time
and place on earth, from the relevant meteorological data’. Although this
calculation was not a crop growth model, it used the hierarchical, explana-
tory approach in that canopy photosynthesis was calculated by integrating
individual leaf photosynthesis over depth in the canopy on the basis of
knowledge of the underlying processes. The integration required the identi-
fication and parameterization of leaf angle distribution functions and the
description of light penetration into the canopy for different conditions of
solar illumination. The estimates of photosynthesis made with this proce-
dure were tested for the potential production situation, and it was found that
the measured rates of canopy photosynthesis could approach the calculated
theoretical values (Alberda, 1968; Alberda & Sibma, 1968).
‘Photosynthesis of leaf canopies’, laid the foundation for the develop-
ment of dynamic models of crop growth. In retrospect, the emphasis on
178 B. A. M. Bouman, H. van Keulen, H. H. van Laar, R. Rabbinge

1965 ‘Photosynthesisof leaf canop~s

-;1

19’10

1975
ARID CROP

1980 ARID CROP

(SAHEL)

PAPRAN

1985

1990

1995

Fig. 3. Pedigree of crop growth simulation models of the “School of de Wit”, 1965-1995.
Models in bold boxes have been “lead” models for the development of other crop models.
Model names are explained in the text.

photosynthesis in crop growth simulation has remained throughout the

years and the models of the ‘School of de Wit’ are all photosynthesis-
driven. One of the first dynamic crop growth simulators was ELCROS
(ELementary CROp Simulator), (de Wit et al., 1970), which was used for
exploratory studies into the potential production levels of crops under
various conditions. This preliminary model contained a detailed, mostly
mechanistic, canopy photosynthesis section, a component describing organ
growth rates and preliminary ideas about crop respiration. Two main-
stream developments in the following years contributed to the evolution of
ELCROS into the first comprehensive model: (i) the quantification of
energy requirements for growth and maintenance processes, both related to
crop respiration; and (ii) the detailed elaboration of crop micrometeorology
in the model MICROWEATHER by Goudriaan (1977). Penning de Vries
et al. (1974) showed that respiration coefficients for growth processes could
be derived, using straightforward stoichiometry, from the biochemical
composition of the biomass. ‘Insight into maintenance processes
was improved’ (Penning de Vries, 197.5) but ‘its quantification remained
 Crop growth simulation models 179

essentially experimental’ (Penning de Vries, 1980). It was only in the early

1990s that more insight into the various processes that contribute to main-
tenance respiration was gained, and that their rates could be theoretically
derived (Bouma, 1995). In MICROWEATHER, crop microweather was
explained as a function of properties of plants and soils, and of the weather
conditions prevalent at some height above the canopy. The elements con-
sidered were solar radiation, energy and mass balances, wind speed and
turbulence. The insights incorporated in this model allowed transpiration
and photosynthesis to be driven by the aerial environment in and above the
crop canopy. The comprehensive model BACROS (BAsic CROp growth
Simulator) was developed from ELCROS (de Wit et al., 1978; Penning de
Vries & van Laar, 1982). BACROS simulates the growth and transpiration
of field crops in the vegetative phase under potential production conditions.
It was designed for grasses such as cereals and specific parameters and
functional relationships specify the actual species under consideration
(Dayan et al., 1981). Although the carbon balance and transpiration are
described mechanistically, partitioning of assimilate and the development
of leaf area are represented empirically. BACROS simulates the growth of
a crop over a whole (vegetative) growing season. For more detailed stu-
dies, the model PHOTON (simulation of daily PHOTOsynthesis and
transpiratioN) was derived from BACROS to simulate photosynthesis.
respiration and transpiration over the course of the day. The time-steps
used in any model should be determined by the smallest time coefficient in
the system. PHOTON therefore uses time-steps of seconds as stomata1
behaviour is considered explicitly. BACROS, on the other hand, uses a
loop to equilibrate this fast process and can thus use a time-step of an
hour without the processes that have much larger time coefficients losing
accuracy and realism. One of the major scientific discoveries using these
comprehensive models was the effect of CO? on stomata1 opening and
hence on photosynthesis and transpiration (de Wit et al., 1978). In these
early years of crop model development, BACROS became the focus for
further development (Fig. 3).

Wuter-limited production situation: ARID CROP

Up until about 1970, the main function of crop models was to explain crop
functioning in a quantitative way, and to explore the potential production
at different geographical locations. One of the first application-oriented
research challenges for modelling was the Dutch/Israeli project ‘Actual and
Potential Production of Semi-Arid Grasslands’ (APPSAG), that was initi-
ated by de Wit in 1970 (Alberda et al., 1992). In this project, crop modelling
was used to quantify and formalize, as far as possible, the relevant processes
involved in water-limited production, and to extrapolate and apply the
180 B. A. M. Bouman, H. van Keulen, H. H. van Laar, R. Rabbinge

resultant knowledge to agricultural production systems (van Keulen et al.,

1982a). ARID CROP (van Keulen, 1975), which was based on the concepts
elaborated in ELCROS and BACROS, was developed to simulate the
growth and water use of fertilized natural pastures in the Mediterranean
region. This comprehensive model successfully coupled a water balance
model, which was later developed into SAHEL (Soils in semi-Arid Habitats
that Easily Leach; Stroosnijder, 1982), to a crop growth model via an
interface between rooting and water uptake. The model describes soil
moisture transport in a simplified way using the unconventional ‘tipping
bucket’ approach, and incorporates a summary sub-model of the carbon
balance and potential canopy transpiration developed from BACROS to
compute the transpiration coefficient of the crop. Potential and actual rates
of crop transpiration were then combined and used to derive the water-
limited crop growth rate. ARID CROP simulates a complete crop growth
cycle from germination, through the stages of vegetative and reproductive
growth and senescence until the death of the crop. However, the processes
of senescence, assimilate partitioning and leaf area development were
represented using relationships that were largely empirical. A revised ver-
sion of the model was shown to conform satisfactorily with experimental
observations for a range of environmental conditions (van Keulen et al.,
1981). ARID CROP was successfully incorporated into an integrated
model of a grazing system comprising separate management and biological
sections, which was used to examine the consequences of contrasting man-
agement strategies in intensive agropastoral systems in a semi-arid region
(Ungar, 1990). ARID CROP was also used in the project ‘Production
Primaire du Sahel’ (PPS) (de Wit, 1975; Penning de Vries & Djiteye, 1982).
The conclusion that production potential was limited in many years by
nutrient deficiency rather than by lack of water (Breman & de Wit, 1983)
was an important outcome of this modelling work in Israel and the Sahel.

The middle years (1980-1990): towards practical applications

During these years, the general emphasis and funding of agricultural

research started to shift from understanding and explaining towards prac-
tical application of the results. Important research issues at that time, such
as agro-ecological zonation, quantitative land evaluation and yield predic-
tion, required exploratory data that were almost impossible to obtain using
conventional methods. The existing comprehensive models, however, were
not very suitable for this purpose because many of the processes were
described in great detail, with a corresponding need for comprehensive
input data, which were often unavailable, and extended computing time.
For production situations where either growth-defining factors alone
 Crop growth simulation models 181

played a role or where water was the limiting factor, knowledge of the
relative importance of the constituent processes allowed the derivation of
effective summary models. For other production situations and levels
however, e.g. where shortage of nitrogen was the limiting factor, there was
still insufficient basic knowledge and the first preliminary and comprehen-
sive models had first to be developed.

Potential and water-limitedproduction situation: SUCROS, WOFOST and

MACROS
SUCROS The first summary model presented was SUCROS (Simple
and Universal CROp growth Simulator; van Keulen et al., 19826).
SUCROS is a simple growth model with a time-step of 1 day, that relies
heavily for its functional relationships on more detailed process-based
models such as BACROS. The original version of SUCROS simulated dry
matter production of a crop from emergence to maturity under potential
production conditions. Like BACROS, SUCROS is universal in nature
because the physical and physiological processes described are applicable
to a wide range of environmental conditions. SUCROS has been applied
to various crops, e.g. wheat, potato and soybean (van Keulen et al.,
1982b) by altering the crop parameters. An updated version, SUCROS87.
was published in 1989, with crop parameters for spring wheat, winter
wheat, maize, potato and sugar beet (Spitters et al., 1989). In 1992, the
latest versions of SUCROS for spring wheat were presented: SUCROSl
for potential production, and SUCROS2 for water-limited production
(van Laar et al., 1992; Goudriaan & van Laar, 1994). In the latter model.
SUCROSl is linked to the soil water balance module SAHEL. Subse-
quently, SUCROS became the lead model for further simplification and
development of specific purpose-oriented models, e.g. INTERCOM
(INTERplant COMpetition) for the interaction between field crops and
weeds (Kropff & van Laar, 1993) and SBjWWFLEVO (Sugar Beet/Winter
Wheat in FLEVOland) which used remotely sensed inputs for growth
monitoring (Bouman, 1992; Fig. 3).

WOFOST One of the first application-oriented models to be derived

from SUCROS was WOFOST (World Food STudies). This model was
developed by the Centre for World Food Studies, an interdisciplinary
group of scientists from the Department of Development Economics of the
Free University of Amsterdam in cooperation with TPE-WAU and
CABO-DLO in Wageningen. The aim of the Centre was to explore the
possibilities of increasing the agricultural productivity of developing
countries (van Keulen & Wolf, 1986; van Diepen et al., 1988). In the
development of the successive versions of WOFOST, the emphasis was on
182 B. A. M. Bouman, H. van Keulen, H. H. van Laar, R. Rabbinge

their practical application for studies on quantitative land evaluation,

regional yield forecasting, analysis of risk and inter-annual yield variation,
and the quantification of the effects of climate change (see Hijmans et al.
(1994) for a review of these WOFOST applications). As in SUCROS, the
process descriptions are universal and the model is tailored to various crops
by altering the crop parameters (van Heemst, 1988). Because of its appli-
cation-orientation, a number of user-friendly features were introduced
in WOFOST. For example, there was a crude geographical information
system facility for presenting the output of the simulations in the form of
maps, and a menu interface allowed the crop type and production situation
and the crop, soil and weather input data files to be selected easily.

MACROS and ORYZA The MACROS modules (Modules of an

Annual CROp Simulator; Penning de Vries et al., 1989) for crops in the
semi-humid tropics were developed as part of the SARP project (Simula-
tion and Systems Analysis for Rice Production). One of the aims of this
project was to transfer the technology of simulation and systems analysis
to multi-disciplinary teams of scientists in Southeast Asia (ten Berge, 1993).
MACROS aided these objectives in two ways: first, as an instructional and
training vehicle for the transfer of agrotechnology and systems analysis;
and second, as a tool for the development and application of models in the
‘cropping systems’, ‘potential production’, ‘water, nutrients and roots’,
and ‘insect pests, diseases and weeds’ research themes. MACROS consists
of a series of basic modules for potential and water-limited crop growth
and for the water balance of soils in both freely draining (SAHEL) and
water-logged conditions (SAWAH; ten Berge et al., 1991). Like SUCROS
and WOFOST, the model is generic and parameters are given for a large
number of crops. Compared with SUCROS, however, MACROS has
retained more of the character of a comprehensive model. An important
feature of MACROS in its role as a training tool, was its transparent,
modular structure that allowed scientists to choose and combine appro-
priate crop growth and water balance modules for addressing their specific
production situations and research questions. Case-studies based on the
MACROS modules were presented at a number of international work-
shops (Penning de Vries et al., 1991) at the end of the second phase of
SARP (1987-1991). In the third and last phase of the project (1992-1995),
the application of the models that had been developed was emphasized by
concentrating research efforts into six application programmes. These
were: (i) agro-ecological zonation and characterization; (ii) optimization of
crop rotations and water use; (iii) application of models in plant breeding
programmes; (iv) evaluation of the impact of climate change on rice pro-
duction; (v) optimization of nitrogen management; and (vi) optimization
 Crop growth simulation models 183

of pest management. A series of rice models based on the MACROS

modules and on SUCROS was developed to serve these specific applica-
tions under the generic name ORYZA (e.g. Kropff et al., 1994).

N-limited production situation: PAPRAN

Nitrogen dynamics in soils and crops were studied under semi-arid
conditions in on-going projects in Israel and the Sahel. However, progress
was slow because the biological and soil chemical processes involved are
complex and difficult to quantify. The first modelling work, which was
based on a relatively simple set of supply and demand functions, resulted in
the description of N uptake and redistribution in plant tissue (Seligman et
al., 1975). Combining these descriptions with those of ARID CROP led to
the development of the preliminary model PAPRAN (Production of Arid
Pastures limited by RAinfall and Nitrogen) for annual pasture production
in semi-arid conditions in which growth is limited by rainfall and nitrogen
(N) (Seligman & van Keulen, 198 1; van Keulen, 1982). PAPRAN is basi-
cally a soil&water balance model where plant growth is closely related both
to the amount of water transpired by the canopy and to its N status, and
where N transformations in the soil are represented by immobilization and
mineralization processes (van Keulen, 1982). Continuation of this line of
work resulted in the development of a comprehensive model for spring
wheat at this production level (van Keulen & Seligman, 1987).

Recent developments (1990-1995): operationalization

The summary models initiated in the 1980s are increasingly being used
operationally as a result of the demand by policy makers and land managers
for data that can only be produced by models. Typical applications include
agro-ecological zonation, regional yield forecasting and scenario studies for
exploring the effect of environmental or socioeconomic changes on agri-
culture. Moreover, the increasing pressure from agricultural funding agen-
cies to ‘prove’ the operational applicability of modelling has led to research
being driven by the development of new technology. This change of
emphasis has introduced new requirements for models and has highlighted
the importance of software quality, an issue that had been recognized in the
early years of modelling, e.g. by Arnold & de Wit (1976).

Operational applications: WOFOST, LINTUL

WOFOST Two major successful applications of WOFOST in the 1990s
were in the policy study ‘Ground for Choices’ (Netherlands Scientific
Council for Government Policy, 1992; Rabbinge & van Latestijn, 1992)
and in the Monitoring Agriculture with Remote Sensing project (MARS)
184 B. A. M. Bouman, H. van Keulen, H. H. van Laar, R. Rabbinge

of the Joint Research Centre (JRC) of the Commission of the European

Communities (Meyer-Roux & Vossen, 1994). In the first study, the model
was used to explore regional yield potentials in the EU under different
management intensities. The results were used to generate technical
coefficients for different crop rotations (de Koning et al., 1995). These
coefficients were then used in the linear programming model GOAL
(General Optimal Allocation of Land use; Scheele, 1992) to optimize land
use and production systems under four contrasting economic scenarios.
The outcomes of these simulations differed widely in terms of the land
required, the costs of production, the employment situation and the use of
fertilisers and pesticides. In the MARS project, WOFOST was integrated
with a geographical information system (GIS) to produce the crop growth
monitoring system (CGMS) for operational yield forecasting for the EU
(van Diepen, 1991; Vossen, 1995). For this purpose, WOFOST was
upgraded to version 6.0 with new routines developed in SUCROS (see
below) and new functionalities developed especially for the MARS project
(Supit et al., 1994). For crop yield forecasting, research is continuing by
exploring possibilities of integrating crop growth models with remote
sensing data to improve the forecasting accuracies (e.g. Bouman, 1995).

LZNTUL For many studies at scales ranging from the regional to the
global, existing summary models needed further simplification because the
availability and quality of model input data were often found to be more
constraining than knowledge of the basic processes incorporated in them.
Spitters (1990) argued that SUCROS could be further simplified by
incorporating only those processes that affect the major determinants of
growth, and laid the foundations for a modelling approach that would
later be baptised LINTUL (Light INTerception and UtiLization; Spitters
& Schapendonk, 1990; Kooman, 1995). LINTUL was the first deviation
from the photosynthesis-based models of the De Wit school. In the
LINTUL models, total dry matter production is calculated using the
Monteith approach (Monteith, 1969, 1990) in which crop growth rate is
calculated as the product of interception of radiation by the canopy and a
light-use efficiency (LUE), which should more correctly be called a dry
matter: radiation quotient (Russell et al., 1989). The LUE can often be
considered constant over the growing season and a property of the crop of
interest. For regional studies, LINTUL-type models have the advantage
that data input requirements are drastically reduced and model para-
meterization is facilitated. The LINTUL approach was used, at the
request of the International Potato Center (CIP), for the agro-ecological
characterization of global potato production to help target research at
production problems in those regions where potato cultivation is most
 Crop growth simulation models 185

promising (van Keulen & Stol, 1995). Recently, Penning de Vries et al.
(1995) used the LINTUL approach in a world food study in which
potential and water-limited food production was estimated for the year
2040 for 15 major regions of the world.

Model quality: SUCROS

SUCROS has been used to test ways of improving model and software
quality. A set of rules and utilities for programming the typical crop growth
models of the ‘School of de Wit’ was developed in FORTRAN and called
the FORTRAN Simulation Environment (FSE; van Kraalingen, 1993).
FSE supports the state-variable approach by organizing process equations
into tasks for initialization, updating of state variables, calculation of rate
variables and calculation of end-of-season characteristics such as harvest
index. These tasks are broadly equivalent to the INITIAL, DYNAMIC
and TERMINAL sections of the early simulation languages (Brennan c,t
al., 1970) with the added clarity of distinguishing between the calculation
of rate and state variables in what would be the DYNAMIC section.
Moreover, the model equations in FSE are separated as much as possible
from the supporting code that takes care of such tasks as data reading, data
checking and output writing. A standard format has been introduced to
take care of input and output data, and modellers using FSE are encour-
aged by its structure to program in an orderly and modular fashion. In
SUCROS87 (Spitters et al., 1989), process descriptions had been organized
into subroutines. Further development has continued in the 1990s and has
resulted in a series of interchangeable, process description routines, e.g. for
light interception, photosynthesis and transpiration, which are universally
applicable but that differ in the level of detail and type of input data needed.
On the basis of the specific model purpose and the availability of input data
(especially relevant in an operational context!), model users can select the
appropriate routines to link to their main model. Finally, standardized
procedures are being developed to test the quality of the software itself, e.g.
to check for programming errors and confirm the reproducibility of results,
and to compare the model outputs with experimental data.

THE CROP MODEL RECORD: USEFULNESS AND

APPLICABILITY

Increasing insight

Crop growth modelling started 30 years ago with the aim of increasing our
insight into crop growth processes by a synthesis of knowledge expressed
186 B. A. M. Bouman, H. van Keulen, H. H. van Laar, R. Rabbinge

using mathematical equations. This is still the main aim of developing

crop growth models. Simulation models are powerful tools for testing our
understanding of crop performance by comparing simulation results and
experimental observations, thus making explicit gaps in our knowledge.
Experiments can then be designed to fill these gaps. This function of crop
growth modelling is difficult to evaluate because it is almost impossible to
predict what would have been our knowledge of crop performance had the
model paradigm not been so pervasive in research (Seligman, 1990). Crit-
ics such as Passioura (1973) and Monteith (1981) have suggested that
complex models cannot show anything that could not be deduced by the
use of straightforward common sense. Crop modellers, however, insist
that no other technique is as powerful for synthesizing knowledge on sub-
processes and increasing our understanding of whole crop behaviour (de
Wit et al., 1978; Loomis et al., 1979; Penning de Vries et al., 1989). Well-
tested crop growth models can be used to explore, in a quantitative way,
the relative importance of crop characteristics, such as physiological and
morphological traits, and environmental characteristics, in a manner that
would not be possible in field experimentation. Sometimes, the simula-
tions produce counterintuitive results. An example of this is found in the
effects of stomata1 behaviour on crop growth and development when the
environment changes (de Wit et al., 1978). These interesting cases stimu-
late further thinking and experimentation and are good evidence for
refuting the criticisms of this type of modelling.

Operational applications

Yield prediction
Validated models can be used in application-oriented research by scientists
and in operational applications where their users are managers or other
non-scientists. Most of the application-oriented research using the crop
growth models of the ‘School of de Wit’ has been related to the problem
of yield prediction (Seligman, 1990) including world food production
studies (Buringh et al., 1979; Penning de Vries et al., 1995), agro-ecologi-
cal zonation (Aggarwal, 1993; van Keulen & Stol, 1995) and explorations
of the effects of climate change on crop production (Wolf, 1993; Matthews
et al., 1995). Operational applications include the use of the crop growth
monitoring system (CGMS) by the Joint Research Centre for producing
monthly yield predictions for the regions of the EU (Vossen, 199.5), and
the use of WOFOST by Dutch consultancy agencies in land use planning
projects (personal communication). Two reasons may be postulated for
the relative success of crop models in yield prediction. First, models are
the only means of systematically exploring the production potential of
 Crop growth simulation model.7 187

agricultural crops in historical or predicted future weather conditions.

Second, there is generally no way of testing the validity of predictions at
regional or global scales. An interesting exception to this second issue was
the recent comparison in the MARS project of yields predicted by CGMS
and by conventional regression techniques with long-term yield statistics
(de Koning et al., 1993). The results indicated that, for most crops, the
accuracy of yield predictions made using only time-series regression mod-
els could not yet be improved by adding the output of the WOFOST
model at the NUTS-O (country) or NUTS1 (primary administrative
region) level. However, because the accuracy of the official yield statistics
is unknown, it was impossible to separate the effects of unrealistic simu-
lations from errors in the statistics. The actual user of CGMS, i.e. the
JRC, however, found many additional benefits of the modelling approach
that were not included in the scientific evaluation. The WOFOST predic-
tions were thus timely(!), objective, quantitative, and consistent over large
areas (Heath, 1991; Vossen (JRC), personal communication). Addition-
ally, the model simulations provided additional means of comparison with
other sources of information, such as field-sampling, remote sensing and
expert knowledge, which are all used to derive the final yield estimates for
the EU. Similar views on the benefits of integrated techniques (including
crop growth models) have been reported by Horie et al. (1992) for rice
yield forecasting in Japan, and Gommes (199 1) for early warning systems
in Africa, Asia and Latin America.

Plant breeding
Crop growth models have been used in plant breeding to simulate the
effects of changes in the morphological and physiological characteristics of
crops and thus to aid in the identification of ideotypes (Donald, 1968) for
different environments (Dingkuhn et al., 1993; Hunt, 1993; Kropff er al.,
1995). Hunt (1993) and Palanisamy et al. (1993) suggested that crop
growth models that have been parameterized for new cultivars in field
experiments can be used to simulate the long-term yield stability of these
cultivars at a location under the expected range of climatic conditions.
This technique holds out the promise of reducing the cost of breeding
programmes by limiting the number and years of expensive, multi-location
trials that are currently required to ensure statistical reliability. A recent
review of literature on the use of modelling in potato breeding led
Ellis&he & Hoogendoorn (1995) to the conclusion that ‘simulation
modelling can contribute to the efficiency of potato breeding programmes,
because modelling analyses complex characteristics, indicates the most
promising components for selection, can forecast plant growth under var-
ious conditions, including biotic and abiotic stress, and helps, therefore.
188 B. A. M. Bouman, H. van Keulen, H. H. van Laar, R. Rabbinge

to formulate breeding strategies’. Breeding objectives can be fine-tuned for

particular regions or environments on the basis of model evaluations
for these circumstances (e.g. Kooman, 1995). The involvement of plant
breeders in recent publications on modelling and breeding suggests a
change in attitude since Seligman’s remark in 1990 that simulation results
using crop growth models have rarely inspired breeders to adapt their
breeding programmes (Seligman, 1990).

Crop management
Crop growth simulation models have been used in numerous studies to
help farmers in day-to-day, i.e. tactical, decision making. They have been
used to investigate the effects of management options such as sowing time,
plant population density, irrigation timing and frequency and fertiliser
applications in different environmental conditions on long-term mean
yield and yield probability (e.g. Ungar, 1990; Carberry et al., 1992, 1993;
Keating et al., 1993; Aggarwal et al., 1994; Aggarwal & Kalra, 1994;
Rotter & Dreiser, 1994). In some cases, these studies have stimulated field
experimentation to test the outcomes predicted in the simulations (ten
Berge et al., 1994). However, the operational application of crop growth
models to support tactical decision making has generally not yet been
successful (Seligman, 1990) with the notable exception of the areas of
irrigation scheduling and water management (van Keulen & Penning de
Vries, 1993) and pest and disease management (e.g. the Epidemics Predic-
tion and Prevention System (EPIPRE); Rabbinge & Rijsdijk, 1983).
Treatment of the latter type of model is outside the scope of this paper and
readers are referred to relevant publications elsewhere, (e.g. Rabbinge et
al., 1990; Kropff & Lotz, 1992; Teng & Savary, 1992).
Recently, researchers have started to apply the results of crop models to
tactical decision making using knowledge based systems such as expert
systems and decision support systems. These software systems have been
promoted since the mid-1980s as a major breakthrough, opening new
horizons in decision support (Schiefer & da Silva, 1995). However, despite
continuing reports of successful developments and prototyping, Hilhorst
& Manders (1995) found that the overall acceptance of the technology in
agriculture in The Netherlands is still limited. They suggested that a main
reason for this lack of acceptance was the knowledge-intensive nature of
such systems. Such systems are of strong interest to research organizations
and this has resulted in: (i) an undue emphasis on problems of a scientific
rather than a practical interest; (ii) poor functionality for non-specialist
users; and (iii) an evolutionary development path which does not accord
with modern software engineering standards. However, these deficiencies,
which might equally well apply to the failure of crop growth models in
 Crop growth simulation mod& 189

operational tactical decision support (see above), are now being addressed.
Three other reasons can explain this particular failure of models. First,
because one of the major sources of yield variation in agriculture is the
variability in weather conditions, success of decision support systems
depends largely on their ability to predict future weather. Even today,
weather predictions are at best reasonably accurate for only a few days
ahead and decision support systems have to rely on probability analyses
using long-term historical or generated weather data (van Keulen &
Penning de Vries, 1993). Second, the lack of accurate input data for soil
and crop characteristics, particularly in respect of their spatial variability,
is often a constraint on successful model applications (van Noordwijk &
Wadman, 1992). Third, models are often used for field conditions whereas
they were developed for rather strictly defined hypothetical production
situations (potential production, water-limited production, etc.) in
uniform fields. In farmers’ fields, several limiting and yield-reducing fat-
tors may occur simultaneously, so that the conditions fall outside the
boundary conditions or domain of validity of the models. This raises the
modeller’s dilemma that for ease of application in a particular practical
situation, models should be as simple as possible and require only a small
number of input data, but that on the other hand, they should be complex
and flexible enough to be able to represent the complex effects of the wide
range of potentially interacting yield-limiting and yield-reducing factors
that might be important for the crop of interest. For situations of poten-
tial production, the summary models developed from the comprehensive
crop growth models of the ‘School of de Wit’ satisfactorily predict crop
behaviour. However, although the processes of photosynthesis and growth
respiration are satisfactorily modelled mechanistically, aspects of main-
tenance respiration and morphogenesis (e.g. organ growth, assimilate
partitioning and leaf area development) are still not well understood and
little progress has been made since the release of BACROS and ARID
CROP. In water-limited conditions, the main effect of water shortage on
the reduction in photosynthesis is well understood and has been satisfac-
torily incorporated in summary models. However, recent experiences in
modelling rice growth in the SARP project have indicated the need for
further study and the inclusion of crop-specific adaptation mechanisms
(Wopereis, 1993).
Major gaps still exist in our knowledge of the effects of nutrient-limita-
tion and it is not yet possible to use mechanistic models directly for farm
level applications (van Keulen & Stol, 199 1; van Keulen & Penning de
Vries, 1993). Therefore the operational use of deterministic models that can
handle the even more complex situations that typify actual farming condi-
tions is still a long way off, and poses new challenges for the years ahead.
190 B. A. M. Bouman, H. van Keulen, H. H. van Laar, R. Rabbinge

One of the ways in which progress is being made in the operationalization

of crop growth models is by using other sources of information, such as
field observations, measurements or remote sensing data, for periodic
adjustment of the state variables. There have been, for example, successes in
the prediction and prevention of epidemics (Rabbinge & Rijsdijk, 1983)
water management and irrigation scheduling (De Falcis et al., 1990; Hill,
1991) and crop growth monitoring (Bouman, 1995). These examples also
illustrate the importance of limited but clear-cut objectives by focusing on
specific problems in tactical decision making.

POSTSCRIPT

This paper deals with a historical overview of crop growth simulation

models from the ‘School of de Wit’. It was, however, not our intention to
suggest that these models are better than other models that have been
developed elsewhere (see Introduction). The authors thank R. S. Loomis
for his helpful comments and suggestions on earlier versions of the
manuscript. G. Russell is thanked for polishing the paper’s English and
suggesting some improvements ‘by the way’.

REFERENCES

Alberda, Th. (1968). Dry matter production and light interception of crop
surfaces. IV. Maximum herbage production as compared with predicted
values. Netherlands Journal of Agricultural Science, 16, 142-53.
Alberda, Th. & Sibma, L. (1968). Dry matter production and light interception of
crop surfaces. III. Actual herbage production in different years as compared
with potential values. Journal of British Grassland Society, 23,206-l 5.
Alberda, Th., Seligman, N. G., van Keulen, H. & de Wit, C. T. (eds) (1992). Food
from dry lands: an integrated approach to planning of agricultural develop-
ment. Kluwer Academic Publishers, Dordrecht, The Netherlands. 211 pp.
Aggarwal, P. K. (1993). Agro-ecological zoning using crop growth simulation
models: characterization of wheat environments in India. In Systems
approaches for sustainable agricultural development, eds F. W. T. Penning de
Vries, P. Teng 8z K. Metselaar. Kluwer Academic Publishers, Dordrecht, The
Netherlands. pp. 97-109.
Aggarwal, P. K. & Kalra, N. (1994). Analyzing the limitations set by climatic
factors, genotype, and water and nitrogen availability on productivity of
wheat II. Climatically potential yields and management strategies. Field
Crops Research, 38, 93-103.
Aggarwal, P. K., Kalra, N., Singh, A. K. & Singha, S. K. (1994). Analyzing the
limitations set by climatic factors, genotype, and water and nitrogen avail-
ability on productivity of wheat I. The model description, parameterization
and validation. Field Crops Research, 38, 73-91.
 Crop growth simulation models 191

Arnold, G. W. & de Wit, C. T. (eds) (1976). Critical evaluation of systems

analysis in ecosystems research and management. Simulation Monographs,
PUDOC, Wageningen, The Netherlands. 108 pp.
Bouma, T. J. (1995). Utilization of respiratory energy in higher plants; require-
ments for ‘maintenance and transport processes’. Ph.D. thesis, Wageningen
Agricultural University. 144 pp.
Bouman, B. A. M. (1992). SBFLEVO and WWFLEVO, Growth models to
simulate crop growth, optical reflectance and radar backscatter of sugar beet
and winter wheat, calibrated for Flevoland. CABO-DLO Report 163.
CABO-DLO, Wageningen, The Netherlands. 61 pp. + appendices.
Bouman, B. A. M. (1994). A framework to deal with uncertainty in soil
and management parameters in crop yield simulation: a case study for rice.
Agricultural Systems, 46, l-17.
Bouman, B. A. M. (1995). Crop modelling and remote sensing for yield
prediction. Netherlands Journal of Agricultural Science, 43, 143361.
Breman, H. & de Wit, C. T. (1983). Rangeland productivity and exploitation in
the Sahel. Science, 221, 1341-7.
Brennan, R. D., de Wit, C. T., Williams, W. A. & Quattrin, E. V. (1970). The
utility of a digital simulation language for ecological modelling. Oecologiu
(Berl.), 4, 113-32.
Buringh, P., van Heemst, H. D. J. & Staring, G. (1979). Potential world food
production. In MOIRA, Model Of International Relations in Agriculture, ed.
H. Linnemann. Contributions to Economic Annals, Vol. 124, North Holland
Publishing Company, Amsterdam/New York, pp. 19-74.
Carberry, P. S., Muchow, R. C., Williams, R., Sturtz, J. D. & McCown, R. L.
A simulation model of kenaf for assisting fibre industry planning in northern
Australia. I. General introduction and phenological model. Australiun
Journal qf Agricultural Research, 43, 150 1-- 15 13.
Carberry, P. S., Muchow, R. C. & McCown, R. L. (1993). A simulation model of
kenaf for assisting fibre industry planning in northern Australia. IV. Analysis of
climatic risk. Australian Journal af’Agricultural Research, 44, 7 13330.
Dayan, E., van Keulen, H. & Dovrat, A. (1981). Experimental evaluation of a crop
growth simulation model. A case study with Rhodes grass. Agro-cco.s~:rtrms.
7, 113--26.
De Falcis, D., Marletto, V. & Zinoni, F. (1990). Irrigation of tomatoes for
industrial processing: reliability of a water balance model. Actrr
Horticulturae, 278, 773-80.
de Koning, G. H. J., van Keulen, H., Rabbinge, R. & Janssen, H. ( 1995).
Determination of input and output coefficients of cropping systems in the
European Community. Agricultural Systems. 48, 4855502.
de Koning, G. H. J., Jansen, M. J. W., Boons-Prins, E. R., van Diepen, C. A. &
Penning de Vries, F. W. T. (1993). Crop growth simulation and statistical
validation for regional yield forecasting across the European Community.
Simulation reports CABO-TT, no. 3 1. CABO-DLO, Wageningen. 105 pp.
de Wit, C. T. (1965). Photosynthesis of leaf canopies. Agricultural Research
Report 663, PUDOC, Wageningen, The Netherlands. 57 pp.
de Wit, C. T. (1970). Dynamic concepts in biology. In Prediction and management
of photosynthetic productivity, Proceedings International Biological Program:
Plant Production Technical Meeting, Trebon, ed. I. Setlik. PUDOC.
Wageningen, The Netherlands. pp. 17723.
192 B. A. M. Bouman, H. van Keulen, H. H. van Laar, R. Rabbinge

de Wit, C. T. (1975). Model studies on actual and potential production in arid

regions. In Proceedings of the Seminar Evaluation and mapping of tropical
African Rangelands, ILCA, Addis Ababa, Ethiopia. pp. 329-3 1.
de Wit, C. T. (1982a). Simulation of living systems. In Simulation ofplunt growth and
cropproduction, eds F. W. T. Penning de Vries & H. H. van Laar. Simulation
Monographs, PUDOC, Wageningen, The Netherlands. pp. 3-8.
de Wit, C. T. (1982b). Coordination of models. In Simulation of plant growth and
crop production, eds F. W. T. Penning de Vries & H. H. van Laar. Simulation
Monographs, PUDOC, Wageningen, The Netherlands. pp. 2631.
de Wit, C. T. & Penning de Vries, F. W. T. (1982). L’Analyse des systemes de
production primaire. In La productivite des pdturages sahe’liens. Une etude des
sols, des vegetations et de l’exploitation de cette ressource naturelle, eds F. W.
T. Penning de Vries & M. A. Djiteye. Agricultural Research Reports 918,
PUDOC, Wageningen, The Netherlands. pp. 20-7.
de Wit, C. T., Brouwer, R. & Penning de Vries, F. W. T. (1970). The simulation
of photosynthetic systems. In Prediction and management of photosynthetic
productivity, Proceedings of the International Biological Program/Plant Pro-
duction Technical Meeting, Trebon, ed. I. Setlik. PUDOC, Wageningen, The
Netherlands. pp. 47-70.
de Wit, C. T. et al. (1978). Simulation of assimilation, respiration and trans-
piration of crops. Simulation Monographs, PUDOC, Wageningen, The
Netherlands. 148 pp.
Dingkuhn, M., Penning de Vries, F. W. T. & Miezan, K. M. (1993). Improve-
ment of rice plant type concepts: systems research enables interaction of
physiology and breeding. In Systems approaches for sustainable agricultural
development, eds F. W. T. Penning de Vries, P. Teng & K. Metselaar. Kluwer
Academic Publishers, Dordrecht, The Netherlands. pp. 19-35.
Donald, C. M. (1968). The breeding of crop ideotypes. Euphytica, 17, 385403.
Ellis&he, D. & Hoogendoorn, J. (1995). The use of models in potato breeding.
In Potato ecology and modelling of crops under conditions limiting growth,
eds A. J. Haverkort & D. K. L. Mackerron. Kluwer Academic Publishers,
Dordrecht, The Netherlands. pp. 341-56.
Gommes, R. A. (1991). The FAO approach to crop monitoring and forecasting. In
Proceedings of the Conference on the Application of Remote Sensing to
Agricultural Statistics, eds F. Toselli & J. Meyer-Roux. Commission of the
European Communities, Luxembourg. pp. 187-97.
Goudriaan, J. (1977). Crop micrometeorology: a simulation study. Simulation
Monographs, PUDOC, Wageningen, The Netherlands. 249 pp.
Goudriaan, J. & van Laar, H. H. (1994). Modelling potential crop growth
processes. Kluwer Academic Publishers, Dordrecht, The Netherlands.
239 pp.
Heath, D. W. (1991). Application of remote sensing to agricultural statistics. In
Proceedings of the Conference on the Application of Remote Sensing to
Agricultural Statistics, eds F. Toselli & J. Meyer-Roux. Commission of the
European Communities, Luxembourg. pp. 19-25.
Hijmans, R. J., Guiking-Lens, I. M. & van Diepen, C. A. (1994). WOFOST, user
guide for the WOFOST 6.0 crop growth simulation model. Technical document
12, DLO Winand Staring Centre, Wageningen, The Netherlands. 145 pp.
Hilhorst, R. & Manders, R. (1995). Critical success and fail factors
for implementation of knowledge based systems at the farm level. In
 Crop growth simulation models 193

A. J. Udink ten Cate, R. Martin-Clouaire, A. A. Dijkhuizen & C. Lokhorst

(eds), 2nd IFAC/IFIP/EurAgEng Workshop on Artifical Intelligence in
Agriculture, IFAC ~ Elsevier Science Ltd. pp. 2977300.
Hill, R. W. (1991). Irrigation scheduling. In Modeling plant and soil sy’stems. eds
J. Hanks & J. T. Ritchie. Agronomy, 31, 491-509.
Horie, T., Yajima, M. & Nakagawa, H. (1992). Yield forecasting. Agricultutoi
Systems, 40, 2 1 l-36.
Hunt, L. A. (1993). Designing improved plant types: a breeder’s viewpoint. In
Systems upproaches for sustainable ugriculturcrl development, eds F. W. T.
Penning de Vries, P. Teng & K. Metselaar. Kluwer Academic Publishers,
Dordrecht, The Netherlands. pp. 3317.
Keating, B. A., McCown, R. L. & Wafula, B. M. (1993). Adjustment of nitrogen
inputs in response to a seasonal forecast in a region of high climatic risk. In
Sy,stems approaches for sustainable agricultural development, eds F. W. T.
Penning de Vries, P. Teng & K. Metselaar. Kluwer Academic Publishers.
Dordrecht, The Netherlands. pp. 233-52.
Klepper, 0. & Rouse, D. I. (1991). A procedure to reduce parameter uncertainty
for complex models by comparison with real system output illustrated on a
potato growth model. Agricultural Systems. 36, 375595.
Kooman. P. L. (1995). Yielding ability of potato crops as influenced
by temperature and daylength. Ph.D. thesis, Wageningen Agricultural
University, Wageningen. 155 pp.
Kropff, M. J. & van Laar, H. H. (eds) (1993). Modelling crop-weed interactions.
CAB International, Wallingford, UK. 274 pp.
Kropff, M. J. & Lotz, L. A. P. (1992). Systems approaches to quantify crop-weed
interactions and their application in weed management. Agricultural Systems,
40, 265-282.
Kropff, M. J., van Laar, H. H. & Matthews, R. (1994). ORYZAI, An
ecophysiological model for irrigated rice production. SARP Research
Proceedings, AB-DLO, Wageningen, The Netherlands. 110 pp.
Kropff, M. J., Haverkort, A. J., Aggarwal, P. K. & Kooman, P. L. (1995). Using
systems approaches to design and evaluate ideotypes for specific environments.
In Eco-regional approaches .for sustainable lund use und food production, eds
J. Bouma, A. Kuyvenhoven, B. A. M. Bouman, J. C. Luyten & H. G. Zandstra.
Kluwer Academic Publishers, Dordrecht, The Netherlands. pp. 4 I7- 35.
Loomis, R. S., Rabbinge, R. & Ng, E. (1979). Explanatory models in crop
physiology. Annual Review of Plant Physiology,, 30, 339967.
Matthews. R. B., Kropff, M. J., Bachelet, D. & van Laar, H. H. (1995). Modelling
the impact of climate change on rice production in Asia. CAB International.
Wallingford, UK. 304 pp.
McCown, R. L., Hammer, G. L., Hargeaves, J. N. G., Holzworth, D. P.
& Freebairn, D. M. (1995). APSIM: a novel software system for model
development, model testing, and simulation in agricultural research.
Agricultural Systems, 50, 255-27 1.
Meyer-Roux, J. & Vossen, P. (1994). The first phase of the MARS project,
(1988)--(1993): overview, methods and results. In Proceedings of’ the
Conjerence on the MARS project: overviewedand perspectives. Commission of
the European Communities, Luxembourg. pp. 33385.
Monteith, J. L. (1969). Light interception and radiative exchange in crop stands. In
Physiological aspects of crop y,ie/d, eds J. D. Easton, F. A. Haskins, C. Y.
194 B. A. M. Bouman, H. van Keulen, H. H. van Laar, R. Rabbinge

Sullivan & C. H. M. van Bavel. American Society of Agronomy, Madison,

Wisconsin. pp. 89-l 11.
Monteith, J. L. (1981). Epilogue: themes and variations. Plant and Soil, 58, 305-9.
Monteith, J. L. (1990). Conservative behaviour in the response of crops to water
and light. In Theoretical Production Ecology: reflection and prospects, eds R.
Rabbinge, J. Goudriaan, H. van Keulen, F. W. T. Penning de Vries & H. H.
van Laar. Simulation Monographs, PUDOC, Wageningen, The Netherlands.
pp. 3-16.
Netherlands Scientific Council for Government Policy (1992). Grounds for
Choices: four perspectives for the rural areas in the European Community,
SDU, The Hague, The Netherlands. 146 pp.
Palanisamy, S., Penning de Vries, F. W. T., Mohandass, S., Thiyagarajan, T. M.
& Kareem, A. A. (1993). Simulation in pre-testing of rice genotypes in Tamil
Nadu. In Systems approaches for sustainable agricultural development, eds F.
W. T. Penning de Vries, P. Teng & K. Metselaar. Kluwer Academic
Publishers, Dordrecht, The Netherlands. pp. 63-75.
Passioura, J. B. (1973). Sense and nonsense in crop simulation. Journal of the
Australian Institute of Agricultural Science, 4, 117-20.
Penning de Vries, F. W. T. (1975). The cost of maintenance processes in plant
cells. Annals of Botany, 39, 77-92.
Penning de Vries, F. W. T. (1980). Simulation models of growth of crops, par-
ticularly under nutrient stress. In Physiological aspects of crop productivity.
Proceedings of the 15th Colloquium, International Potash Institute, Bern.
pp. 213-26.
Penning de Vries, F. W. T. & van Laar, H. H. (1982). Simulation of
growth processes and the model BACROS. In Simulation of plant growth and
crop production, eds F. W. T. Penning de Vries & H. H. van Laar. Simulation
Monographs, PUDOC, Wageningen, The Netherlands. pp. 114-36.
Penning de Vries, F. W. T. & Djiteye, M. A. (eds) (1982). La productivite des
paturages sahelien; Une etude des sols, des vegetations et de l’exploitation de
cette ressource naturelle. Agricultural Research Reports 918, PUDOC,
Wageningen, The Netherlands. 525 pp.
Penning de Vries, F. W. T., Brunsting, A. B. & van Laar, H. H. (1974). Products,
requirements and efficiency of biological synthesis, a quantitative approach.
Journal of Theoretical Biology, 45, 339-77.
Penning de Vries, F. W. T., van Laar, H. H. & Kropff, M. J. (eds) (1991). Simula-
tion and systems analysis for rice production (SARP). PUDOC, Wageningen,
The Netherlands. 369 pp.
Penning de Vries, F. W. T., van Keulen, H. & Rabbinge, R. (1995). Natural
resources and limits of food production in 2040. In Eco-regional approaches
for sustainable land use and food production, eds J. Bouma, A. Kuyvenhoven,
B. A. M. Bouman, J. C. Luyten & H. G. Zandstra. Kluwer Academic Pub-
lishers, Dordrecht, The Netherlands. pp. 65-87.
Penning de Vries, F. W. T., Jansen, D. M., ten Berge, H. F. M. & Bakema, A.
(1989). Simulation of ecophysiological processes of growth in several annual
crops. Simulation Monographs, PUDOC, Wageningen, The Netherlands.
271 pp.
Plentinger, M. C. & Penning de Vries, F. W. T. (1995). CAMASE, Register of
Agro-ecosystems Models, version 1. AB-DLO, Wageningen, The Netherlands.
179 pp.
 Crop growth simulation models 195

Rabbinge, R. (1986). The bridge function of crop ecology. Netherlands Journal d

Agricultural Science, 3, 239-51.
Rabbinge, R. (1993). The ecological background in food production. In Crop
protection and sustainable agriculture. John Wiley & Sons, Chichester (Ciba
Foundation Symposium 177), pp. 2-29.
Rathinge, R. & Rijsdijk, F. H. (1983). EPIPRE: A disease and pest management
system for winter wheat, taking account of micrometeorological factors.
EPPO Bulletin, 13, 297-305.
Rabbinge, R. & de Wit, C. T. (1989). Systems, models and simulation. In Simu-
lation and systems management in crop protection, eds R. Rabbinge, S. A.
Ward & H. H. van Laar. Simulation Monographs, PUDOC, Wageningen.
The Netherlands. pp. 3-l 5.
Rabbinge, R. & van Latestijn, H. C. (1992). Long-term options for land use in
the European Community. Agricultural Systems, 40, 195-210.
Rabbinge, R., Rossing, W. A. H. & van der Werf, W. (1990). The bridge
function of production ecology in pest and disease management. In Thro-
retical Production Ecology.. reji’ection and prospects, eds R. Rabbinge,
J. Goudriaan, H. van Keulen, F. W. T. Penning de Vries & H. H. van Laar.
Simulation Monographs, PUDOC, Wageningen, The Netherlands. pp.
181-95.
RGtter, R. & Dreiser, C. (1994). Extrapolation of maize fertiliser trial results by
using crop-growth simulation: results for Murang’a District, Kenya. In The
future of the land: mobilising and integrating knowledge for land use options.
eds L. 0. Fresco, L. Stroosnijder, J. Bouma & H. van Keulen. John Wiley &
Sons Ltd., West Sussex, UK. pp. 249-60.
Rossing, W. A. H., Daamen, R. A. & Jansen. M. J. W. (1994). Uncertainty
analysis applied to supervised control of aphids and brown rust in winter
wheat. Part I. Quantification of uncertainty in cost-benefit calculations.
Agricultural Systems, 44, 4 19-48.
Russell, G., Jarvis, P. G. & Monteith, J. L. (1989). Absorption of solar radiation
and stand growth. In Plant canopies, their growth form andfunction. eds G.
Russell, P. G. Jarvis & B. Marshall. Cambridge University Press. Cambridge.
UK. pp. 21-39.
Scheele, D. (1992). Formulation and characteristics of GOAL. Technical Working
Document (W64), Netherlands Scientific Council for Government Policy. The
Hague. 64 pp.
Schiefer, G. W. & da Silva, C. A. (1995). An evaluation of expert systems tech-
nology for decision support. In 2nd IFAC/IFIP/EurAgEng Workshop on
Artificial Intelligence in Agriculture, eds A. J. Udink ten Cate. R. Martin-
Clouaire, A. A. Dijkhuizen & C. Lokhorst. IFAC - Elsevier Science Ltd.
pp. 301-7.
Seligman, N. G. (1990). The crop model record: promise or poor show’! In
Theoretical Production Ecology: rqflection and prospects, eds R. Rabbinge. J.
Goudriaan, H. van Keulen, F. W. T. Penning de Vries & H. H. van Laar.
Simulation Monographs, PUDOC, Wageningen, The Netherlands. pp.
249-64.
Seligman, N. G. & van Keulen, H. (1981). PAPRAN: A simulation model of
annual pasture production limited by rainfall and nitrogen. In Simulution of
nitrogen behaviour of soil-plant systems, eds M. J. Frissel & J. A. van Veen.
PUDOC, Wageningen. pp. 192-220.
196 B. A. M. Bouman, H. van Keulen, H. H. van Laar, R. Rabbinge

Seligman, N. G., van Keulen, H. & Goudriaan, J. (1975).

. , An elementarvd model
of nitrogen uptake and redistribution by annual plant species. Oecologia
(Berl.), 21, 243-61.
Sinclair, T. R., Goudriaan, J. & de Wit, C. T. (1977). Mesophyll resistance and
CO2 compensation concentration in leaf photosynthesis models.
Photosynthetica, 11, 56-65.
Spitters, C. J. T. (1990). Crop growth models: their usefulness and limitations.
Acta Horticulturae, 267, 345-63.
Spitters, C. J. T. & Schapendonk, A.H.C.M. (1990). Evaluation of breeding
strategies for drought tolerance in potato by means of crop growth
simulation. Plant and Soil, 123, 193-203.
Spitters, C. J. T., van Keulen, H. & van Kraalingen, D. W. G. (1989). A simple
and universal crop growth simulator: SUCROS87. In Simulation and sys-
tems management in crop protection, eds R. Rabbinge, S. A. Ward & H. H.
van Laar. Simulation Monographs, PUDOC, Wageningen, The Netherlands.
pp. 147-81.
Stroosnijder, L. (1982). Simulation of the soil water balance. In Simulation of
plant growth and crop production, eds F. W. T. Penning de Vries & H. H. van
Laar. Simulation Monographs, PUDOC, Wageningen, The Netherlands. pp.
175-93.
Supit. I.. Hooiier. A. A. & van Dienen. C. A. feds) (1994). Svstem description of
the WOF&T 6.0 crop growth simulation model implemented in ‘CGMS.
Joint Research Centre, Commission of the European Communities, Brussels,
Luxembourg. 144 pp.
ten Berge, H. F. M. (1993). Building capacity for systems research at national
agricultural research centres: SARP’s experience. In Systems approaches for
agricultural development, eds F. W. T. Penning de Vries, P. Teng &
K. Metselaar. Kluwer Academic Publishers, Dordrecht, The Netherlands.
pp. 515-38.
ten Berge, H. F. M., Jansen, D. M., Rappoldt, C. & Stol, W. (1991). The
soil water balance module SAWAH: User’s guide and outline. Simulation
Report CABO-TT 22. CAB@DLO, Wageningen, The Netherlands. 78 pp.
+ appendices.
ten Berge, H. F. M., Wopereis, M. C. S., Riethoven, J. J. M., Thiyagarajan, T.
M. & Sivasamy, R. (1994). The ORYZA_0 model applied to optimize
nitrogen use in rice. In Nitrogen economy of irrigated rice: field and
simulation studies, eds H. F. M. ten Berge, M. C. S. Wopereis & J. C. Shin.
SARP Research Proceedings, AB-DLO, Wageningen, The Netherlands. pp.
235-53.
Teng, P. S. & Savary, S. (1992). Implementing the systems approach in pest
management. Agricultural Systems, 40, 237-64.
Tsuji, G. Y., Uehara, G. & Bala, S. (eds) (1994). DSSAT Version 3.0. DSSAT
approach to input management. University of Hawaii, Honolulu, Hawaii,
USA.
Uehara, G. & Tsuji, G. Y. (1993). The IBSNAT project. In Systems approaches
jar sustainable agricultural development, eds F. W. T. Penning de Vries,
P. Teng & K. Metselaar. Kluwer Academic Publishers, Dordrecht, The
Netherlands. pp. 505-13.
Ungar, E. D. (1990). Management of agropastoral systems in a semiarid region.
Simulation Monographs, PUDOC, Wageningen, The Netherlands. 221 pp.
 Crop growth simulation models 197

van Diepen, C. A. (1991). An agrometeorological model to monitor the crop

state on a regional scale in the European Community: concept, imple-
mentation and first operational outputs. In Proceedings of the Conference on
the application of remote sensing to agricultural statistics, eds Toselli &
Meyer-Roux. Commission of the European Community, Luxembourg. pp.
269-79.
van Diepen, C. A., Rappoldt, C., Wolf, J. & van Keulen, H. (1988). Crop growth
simulation model WOFOST. Documentation Version 4. I. CWFS, Amsterdam
- Wageningen. 299 pp.
van Duivenbooden, N. & Gosseye, P. A. (eds) (1990). Competition pour des
ressources limit&es: le cas de la cinquiime rtgion du Mali. Rapport 2. Produc-
tions v&g&tales, animal et halieutiques. CABO, Wageningen, The Netherlands.
266 pp.
van Heemst. H. D. J. (1988). Plant data values required for simple crop growth
simulation models: review and bibliography. Simulation Report CABO--TT
17, CABO, Wageningen, The Netherlands. 100 pp.
van Keulen, H. (1975). Simulation of water use and herbage growth in
arid vregions. Simulation Monographs. PUDOC. Wageningen, The
Netherlands. 176 pp.
van Keulen, H. (1982). Crop production under semi-arid conditions. as deter-
mined by nitrogen and moisture availability. In Sirnulution of’pplunt g,ol\,th
and crop production, eds F. W. T. Penning de Vries & H. H. van Laar.
Simulation Monographs, PUDOC, Wageningen. The Netherlands. pp.
234-5 I.
van Keulen, H. & Wolf, J. (eds) (1986). Modelling of agricultural production:
weather, soils and crops. Simulation Monographs. PUDOC. Wageningen.
The Netherlands. 479 pp.
van Keulen, H. & Seligman, N. G. (1987). Simulation of water use. nitrogen
nutrition and growth of a spring wheat crop. Simulation Monographs,
PUDOC, Wageningen, The Netherlands. 310 pp.
van Keulen, H. & Stol, W. (1991). Quantitative aspects of nitrogen nutrition in
crops. Fertilizer Research, 27, 15 l-60.
van Keulen, H. & Penning de Vries, F. W. T. (1993). Farming under uncertainty:
terminology and techniques. In International Crop Science I. Crop Science
Society of America, 677 S. Segoe Rd., Madison, USA. pp. 13944.
van Keulen, H. & Stol, W. (1995). Agro-ecological zonation for potato produc-
tion. In Potato ecology und rnodelling qf’ crops under conditions limiting
grort,th, eds A. J. Haverkort SC D. K. L. Mackerron. Kluwer Academic
Publishers, Dordrecht, The Netherlands. pp. 357- 7 I.
van Keulen, H., Seligman, N. G. & Benjamin, R. W. (1981). Simulation of water
use and herbage growth in arid regions a re-evaluation and further
development of the model ‘Arid crop’. Agricultural Systems, 6. 159 93.
van Keulen, H., Benjamin, R. W., Seligman, N. G. & Noy-Meir, 1. (1982n). Final
technical report and annotated bibliography, Joint Dutch -Israeli project
Actual and potential production from semi-arid grasslands Phase II.
CABO, Wageningen, The Netherlands. 61 pp.
van Keulen, H., Penning de Vries, F. W. T. & Drees, E. M. (I 982h). A summary
model for crop growth. In Simulation ofplant grolt.th and crop production. eds
F. W. T. Penning de Vries & H. H. van Laar. Simulation Monographs.
PUDOC, Wageningen. The Netherlands. pp. X7 -98.
198 B. A. M. Bouman, H. van Keulen, H. H. van Laar, R. Rabbinge

van Kraalingen, D. W. G. (1993). The FORTRAN Simulation Environment

(FSE). In Modelling crop-weed interactions, eds M. J. Kropff & H. H. van
Laar. CAB International, Wallingford, UK. pp. 187-200.
van Laar, H. H., Goudriaan, J. & van Keulen, H. (eds) (1992). Simulation of crop
growth for potential and water-limited production situations (as applied to
spring wheat). Simulation Report CABO-TT, 27, CABO-DLO, Wageningen,
The Netherlands. 105 pp.
van Noordwijk, M. & Wadman, W. P. (1992). Effects of spatial variability of
nitrogen supply on environmentally acceptable nitrogen fertilizer application
rates to arable crops. Netherlands Journal of Agricultural Science, 40, 51-72.
Vossen, P. (1995). Early assessment of national crop yields: the approach devel-
oped by the MARS-STAT Project on behalf of the European Commission.
In Pre-proceedings Seminar ‘Yield Forecasting’, 24-27 October 1995,
Villefranche-sur-Mer. pp. 32347.
Wolf, J. (1993). Effects of climate change on wheat production potential in the
European Community. European Journal of Agronomy, 2,28 l-292.
Wopereis, M. C. S. (1993). Quantifying the impact of soil and climate variability
on rainfed rice production. Ph.D. thesis, Wageningen Agricultural
University, Wageningen. 188 pp.