Annals of Botany 88: 279±291, 2001

doi:10.1006/anbo.2001.1458, available online at http://www.idealibrary.com on

Dynamic Model for the E€ects of Soil P and Fertilizer P on Crop Growth, P Uptake and
Soil P in Arable Cropping: Model Description
D . J . G R EE N WO O D *{, T. V. K A R P I N E T S{ and D . A . S TO N E {
{Horticulture Research International, Wellesbourne, Warwick, CV 35 9EF and {The Institute for the Protection of
Soil from Erosion, Karl Marx Street, 70, 6, Kursk, Russia

Received: 24 November 2000 Returned for revision: 29 January 2001 Accepted: 10 April 2001

A mechanistic model is described for calculating the e€ects of starter fertilizer, granular fertilizer and 0
5 M NaHCO3
pH 8
5 extractable soil phosphate on plant growth and plant P concentration during the entire period of growth up to
commercial harvest. For each day, the model calculates the increment in root growth and partitions it into segments
between the regions of soil enriched with starter fertilizer, those enriched with granular fertilizer and the remainder of
the soil. It calculates the maximum possible amount of P that can di€use through soil to each root segment in each
region. Using this information and the P concentration in the plant, total P uptake is calculated. The increment in
plant weight and root growth is calculated from the current plant weight, plant P concentration and air temperature.
Subroutines calculate daily soil water content, the extractable and non-extractable soil P, and di€usion coecients in
the P-depleted zones around each root segment and in the remainder of the soil. Model simulations and sensitivity
analyses indicate that extractable soil P and starter fertilizer P can lead to higher crop yields than are achieved when
granular fertilizers are incorporated in soil, in the usual way, immediately before sowing. They also indicate that in the
long-term, levels of extractable soil P will move towards a level characteristic of the soil. These ®ndings are in
agreement with results of long-term ®eld experiments that have been reported in the literature. All inputs to the model
that have a substantial impact on P-response of a single crop are easy to obtain. They include standard soil properties,
the maximum potential yield, and daily rainfall, mean air temperature and evaporation from an open water surface.
The model runs interactively at: www.qpais.co.uk/phosmod/phos.htm # 2001 Annals of Botany Company

Key words: Simulation, dynamic model, vegetable crops, soil phosphate, phosphate fertilizer, growth, response, plant
composition.

I N T RO D U C T I O N that they seldom attempt to take account of the enormous

Phosphate de®ciency limits yields in much of the world,
especially in poorer countries where fertilizers are una€ord-
able. On the other hand, in some richer countries, levels of
soil phosphate have often become far higher than are
needed by crops and are causing environmental pollution.
There is, however, still much uncertainty about how best to
adjust fertilizer and manurial practices for di€erent crops
and soil conditions. The growing interest in sustainable
agriculture and horticulture is an incentive to provide better
methods of optimizing practices.
Crop response to soil phosphate depends on a range of
interacting soil and plant processes, many of which have
been studied in detail. One way of applying this information
for predictive purposes is to devise mechanistic models in
which key processes are represented by equations that are re-
calculated for each time interval during the growing season.
Models for P response have been reviewed by Amijee et al.
(1991), Rengel (1993), Claassen and Steingrobe (1999) and
Tinker and Nye (2000). However, while much attention has
been devoted to the soil aspects that a€ect P response, plant
aspects have been largely neglected. Thus, there has been
little attention to the dependence of uptake on plant weight
or plant composition. Another de®ciency of the models is
* For Correspondence. E-mail duncan.greenwood@hri.ac.uk
0305-7364/01/080279+13 $35.00/00
spatial variation of phosphate concentration in soil resulting
from the application of starter and granular fertilizers. This
results in parts of the root system exploring high-P soil
whereas the remainder explores low-P soil. Of the P-
response models so far developed, most attention has been
given to the Barber-Cushman model (Barber and Cushman,
1981; Barber, 1995) for calculating plant uptake of P. It is
based on an equation for di€usive/mass ¯ow of phosphate
through the soil to cylindrical roots. The ®nite di€erence
form of this equation is solved by the Crank-Nickolson
procedure with boundary conditions de®ned by assuming
that root P uptake is related to P concentration at the root
surface by a Michaelis-Menten equation. Root growth is
simulated using a predetermined relationship between the
rate of root elongation and time. Good agreement has been
reported between simulated and measured P uptakes
especially in pot experiments (e.g. Bidin and Barber, 1985;
Jungk and Claassen, 1989) while agreement is poor in some
®eld experiments (Silverbush and Barber, 1984; Lu and
Miller, 1994).
The fundamental drawback of this type of model is that it
does not allow inputs that vary with time (Yanai, 1994).
For example, the soil water content and the pattern of root
growth remain constant and are una€ected by weather or
plant growth. This limitation severely restricts the useful-
ness of the model.
# 2001 Annals of Botany Company
280 Greenwood et al.ÐDescription of Model for Crop Response to Phosphate
More simple, stationary state models have been devised,
including one by van Noordwijk et al. (1990) which has
given promising predictions of the extractable soil P
required for maximal yields of di€erent crops grown on a
range of soils. It is, however, based on a stationary state
concept of the various processes and so does not allow
account to be taken of factors such as changes in daily
weather during growth.
Other modelling approaches have been adopted that do
not su€er from the aforementioned constraints. They have
been widely used in the development of plant growth
models (e.g. Thornley and Johnson, 1990) and in modelling
N-nutrition of plants (e.g. Greenwood et al., 1996; Smith et
al., 1996). They allow the rate and state variables to be
updated for each day, or for shorter intervals, in response to
changes in environmental conditions and crop develop-
ment. An advantage of this approach is that the models can
be readily incorporated into broader models and decision
support systems such as those referred to by Hartkamp
et al., (1999). The purpose of our work was to explore the
possibility of developing a dynamic model along these lines
to explain the P response of vegetable crops. It was hoped
that such a model would allow prediction of the time course
of the response of both plant weight and plant P
concentration, rather than just P uptake, to increasing
amounts of soil P and fertilizer P, an objective that has not,
as far as we are aware, been met by any previous model.
An approach to this problem was suggested by a model
on the related topic of crop response to soil potassium
(Greenwood and Karpinets, 1997a,b). It included algo-
rithms for the dependence of K uptake from the soil on
plant demand and composition. Plant demand for P is
much smaller than that for K, and the P ®xation and
adsorption phenomena are very di€erent from the corre-
sponding K processes. Transport of P through soil is an
order of magnitude slower than that for K. While it is a
reasonable approximation to consider that K is uniformly
distributed through soil, as in the K model, we do not
believe that a similar assumption is tenable for P models.
The intention was therefore to use the K model as a starting
point for the P model and, in particular, to treat P uptake
from regions enriched with fertilizer separately from that in
unenriched regions.
MODEL
Overview
The model simulates the e€ects of soil P, granular fertilizer
P, and starter fertilizer P, on daily crop growth, on P
concentrations within the plant and on the P economy of
the soil. It is considered that granular and starter fertilizers
are applied immediately before sowing, whereas the soil P is
the net result of the soil's type and history. All other
nutrients, including N and K, are considered to be sucient
for maximum growth. The model consists of equations
representing the more important processes and requires
only inputs for which most values can be readily obtained.
Integration is in steps of 1 d as most weather data are only
available as a daily average.
The soil is visualized as being homogeneous with all roots
con®ned to the plough layer, 30 cm in depth. This approxi-
mation is made because nearly all the roots (and thus P
absorption by them) occur in that layer. P is assumed to
exist as extractable soil P (with 0
5 M NaHCO3 pH 8
5),
solution soil P and non-extractable soil P, that includes all
the remaining P. Transport of P through soil is by di€usion
and it is assumed that no P is leached below the plough
layer.
A simpli®ed ¯ow diagram for the P response of a crop
grown on unfertilized soil is given in Fig. 1. The main soil
processes are on the left-hand side of the diagram and the
main plant ones on the right-hand side. The model is,
however, more complex than is implied by the diagram.
There are three regions of soil: one consists of soil enriched
by starter fertilizer, another of soil enriched with granular
fertilizer, and another consists of unenriched soil. The
processes in each region are treated separately. The daily
increment in root length is partitioned into segments that
are allocated to one or more regions. Around each root
segment a zone of depleted soil P develops, which increases
with time throughout the growing period. The soil
processes given on the left of the diagram are re-calculated
for each depletion zone for each day during growth. They
are also re-calculated for the undepleted regions of soil as
there is generally interchange between the extractable and
non-extractable P. The sum of the total amounts of P that
can be transported to each of the root segments gives the
maximum possible uptake. This is, however, reduced
depending on the plant % P to give the actual uptake.
All the above calculations are adjusted for daily rainfall,
temperature and evaporation from an open water surface.
For each day the following are calculated:
(1) the increment in dry weight and the new value of total
plant weight (calculated from the % P in the plant, the
dry weight of the plant and the mean air temperature);
(2) the increment in root length and the total root length
(calculated from the increment in plant weight and
total plant weight) and the partitioning of root growth
into segments that enter regions of soil enriched with
the starter or the granular fertilizer or unenriched
regions;
(3) P uptake calculated from P di€usion to each of the root
segments and the plant % P;
(4) the average volumetric water content of the soil, the
values of the tortuosity coecients, the P-di€usion
coecients for the bulk soil and for each of the P
depletion zones around each segment;
(5) the volume of the P depletion zones around each of the
root segments; the changes in the various amounts of
solution P, extractable soil P and non-extractable soil P
in each of the zones and in the undepleted soil.
Equations
The more important equations and rules are now described.
Symbols and default values are summarized in Tables 1
and 2.
 Greenwood et al.ÐDescription of Model for Crop Response to Phosphate 281

by R. eqn (1)
Daily weather Daily
Temperature ( T )
temp increment in
and volumetric water content plant dry
( V)
1 matter Plant dry
( D W) matter
eqns (1) and (2) ( W)
by eqn (12)
W=W+DW
by G temp

Daily

increment in

root length

( DLt)
Diffusion
eqn (3)
coefficient of P P conc in

through soil plant

D s

eqns(11),(14) and (16) P= W/Up

Root length Lt
by eqn (12)
Lt=Lt+DLt

by eqn (11)

by G . eqn (8)
f
Amount Amount

of non- of

extractable extractable Flux of P

P in P in to root Actual P uptake

upper 30 upper 30 surface Up
cm of cm of eqns eqn (8)
soil soil (6),(7) and (9)

SOIL PROCESSES PLANT PROCESSES

F I G . 1. Simpli®ed ¯ow diagram of the model; details of equations are given in the body of the text.

Daily increment in plant dry matter. The daily increment
in plant weight is given by
DW ˆ Gwt Gtemp d† R d† W d†=‰W0:5 ‡ W d†Š
with minor modi®cations to improve the integration
procedure. W is plant dry weight excluding ®brous roots
which must be expressed in t ha ÿ1, W0
5 is a constant that is
always set at 1 t ha ÿ1, Gwt is a growth coecient, Gtemp(d)
corrects for the e€ects of air temperature on growth and
R(d) for % P; the d in parentheses indicates that the
variable depends on the day from Jan 1. The integral of this
equation with Gtemp(d) and R(d) both set equal to one has
given good ®ts to measurements of W made during the
1† main growing season in western European ®eld experiments
(e.g. Greenwood et al., 1977, 1991).
Gtemp d† ˆ 1 for 12 5 Atemp 4 30  C
Gtemp d† ˆ Atemp ÿ 7†= 12 ÿ 7† for 7 5 Atemp 4 12  C
Gtemp d† ˆ 0:01 for Atemp 4 7  C
where Atemp is the mean daily air temperature.
282 Greenwood et al.ÐDescription of Model for Crop Response to Phosphate
T A B L E 1. De®nition of symbols and default values

Symbol De®nition Default value

Agfert Weight of granular fertilizer P incorporated in soil before sowing 0
05 t ha ÿ1

Astfert Weight of starter fertilizer P incorporated in soil before sowing 0
01 t ha ÿ1
a Root radius 0
008 cm
Bd Bulk density 1
4 g cm ÿ3
Dw Di€usion coecient of P through water at 25 8C 1
02 cm2 d ÿ1
Gafr Coecient in Freundlich isotherm 51
7
Gbfr Coecient in Freundlich isotherm 0
505
Gp Parameter for e€ect of P concentration in the plant dry matter on P uptake by the roots 0
6
Kcn Coecient for the decline in plant P parameters with increase in plant weight 0
0209 t ÿ1 ha
Kextr Rate constant for conversion of extractable P into non-extractable soil P 6
37  10 ÿ4 d ÿ1
Knextr Rate constant for conversion of non-extractable soil P into extractable soil P 3
32  10 ÿ5 d ÿ1
Phlf(0) Parameter used in de®ning e€ect of % P in plant dry matter on growth rate; value when W ˆ 0 0
04 %
Pextrw Concentration of extractable P in upper 30 cm of soil expressed per unit weight of soil 18 mg g ÿ1
Pnextrw Concentration of non-extractable P in upper 30 cm of soil 1190 mg g ÿ1
Pmax(0) The minimum % P of dry matter required for maximum growth rate when W ˆ 0 0
6 %
Pmin Minimum possible % P of plant dry matter 0
1 %
Popt(0) Parameter for e€ect of % P of plant dry matter on the rate of plant P uptake when W ˆ 0 0
3 %
Pseed The % P of seed dry matter 0
8 %
Rgrno Radius of P fertilizer granule 1
7 mm
Sdef Soil moisture de®cit on Jan 1st 0 mm
Th Time from sowing to harvest 146 d
Ts Sowing date in days from Jan 1st 125 d
Vfc Volumetric water content at ®eld capacity 0
26
Vlsens Volumetric water content used in sensitivity analyses 0
2
Vsoil Volume of soil to depth of 30 cm 3  103 m3
Vst Volume of soil enriched by starter fertilizer 24 m3
Wmax Plant dry matter yield at harvest 14
5 t ha ÿ1
W0 Dry weight of sown seed 1
64 kg ha ÿ1
W0
5 Constant relating growth rate to W 1
0 t ha ÿ1

P concentration in the plant dry matter and weights of plant dry matter are for the entire plant excluding ®brous roots.

T A B L E 2. De®nition of symbols without default values

Symbol De®nition Units

(d) Indicates that the variable depends on day from Jan 1 d

Droot Depth of soil containing 90 % of roots cm
D1 Di€usion coecient of P through water after correction for temperature cm2 d ÿ1
Ds Di€usion coecient of P through soil cm2 d ÿ1
Fp Flux of P to root surface mg cmÿ2 d ÿ1
Ftort Tortuosity in calculation of di€usion coecient of P through soil Dimensionless
Gf Actual increment of plant P uptake expressed as a fraction of the maximum Dimensionless
Gtemp Corrects daily growth rate for mean air temperature Dimensionless
Gwt Growth rate coecient t ha ÿ1 d ÿ1
Lt Total root length km m ÿ2
nLdpl Length of root segment cm
F Viscosity of water cP
Pgrth P concentration in the plant dry matter on any day during growth %
Pl Concentration of P in soil solution mg cmÿ3
Pextrv Concentration of extractable soil P expressed as per unit volume of soil i.e. Pextrv ˆ Bd Pextrw mg cmÿ3
Pextrvo Pextrv in regions of soil una€ected by root uptake mg cmÿ3
R Factor for reducing the increment in growth for sub-optimal % P Dimensionless
Rgrn Radius of the enriched zone around a fertilizer granule mm
Tdf Time from formation of a given root increment d
Tg Time from incorporating granules in soil d
Ttemp Temperature 8C
Up Crop uptake of P kg ha ÿ1
Vdpl Volume of the depleted zone round each root segment cm ÿ3
V1 Volumetric water content of soil on a given day cm3 cmÿ3
W Total plant dry weight excluding ®brous roots t ha ÿ1
 Greenwood et al.ÐDescription of Model for Crop Response to Phosphate
These relationships were based on experience of growing
vegetable crops in the UK but are similar to those derived
from measurements made on cauli¯ower by Alt et al.,
(2000). They may, however, vary depending on the cultivar
and environment.
R d† ˆ minf1; ‰ 1 ‡ P hlf 0 W†=P max 0 W†††=
2†
1 ‡ P hlf 0 W†=P grth 0 d†††Šg
where
P hlf 0 W† ˆ Phlf 0† 1 ÿ Kcn W†
P grth 0 ˆ Pgrth ÿ Pmin
P max 0 W† ˆ Pmax 0† 1 ÿ Kcn W† ÿ Pmin
where Pgrth is the % P in the plant at time t (days), Pmin is
the minimum % P that can occur in the plant, Pmax is the
lowest % P in the plant at which the growth rate is
maximal; and Phlf is a coecient de®ning the sensitivity of
R(d) to % P in the plant. The zero in parentheses indicates
values of the parameters when W is zero; they decline
linearly with an increase in W; the coecient for this
decline is given by Kcn. Evidence for this decline is provided
by Belanger and Richards (1999) amongst others. Equation
(2) is derived by modifying the Michaelis-Menten equation
to ensure that R(d) ˆ 1 when P grth 0 (d) reaches P max 0 (W);
it also ensures that R(d) does not continue to increase with
further increase of P grth 0 (d) beyond P max 0 (W).
Daily increment in root growth. The daily increment in
root growth is calculated as
DLt ˆ Al ‰e A1 ln W d††‡Bl †
ŠDW d†=W d† 3†
where W is plant weight in t ha ÿ1, Lt is root length in km
m ÿ2 and Al and Bl are coecients. The equation is derived
by di€erentiating:
ln Lt ˆ Al ln W ‡ Bl 4†
For non-leguminous vegetable crops grown on sandy loam
soils,
Al ˆ 1
07; Bl ˆ 1
81 for ln W 4 ÿ 3
Al ˆ 0
82; Bl ˆ 1
06 for ln W 4 ÿ 3
Equation (4), with the indicated parameter values, ®tted
experimental data for a wide range of crops (Greenwood et
al., 1982; Costigan, 1985; Bosch Serra et al., 1997). There is
also evidence that root growth may be similar over a wide
range of soil conditions (e.g. Stypta et al., 1987). When ln
W 4 ÿ 3 the equations above may be widely applicable.
Root growth in the model is not enhanced by plant P or
plant water stress.
Partitioning of the roots. All root absorption of P is
considered to take place from the upper 30 cm of soil.
Throughout growth, a fraction of the total root length
enters regions of soil enriched with granular fertilizer. This
fraction, which is small, equals the ratio of the volume
283
occupied by the regions of soil enriched with granular
fertilizer divided by the total volume of soil to a depth of
30 cm. The rest of the root system initially enters the region
of soil enriched with starter fertilizer that occupies 0
8 % of
the total volume of soil. However, root growth in this
region ceases when the volume of the di€usion zones
calculated using eqn (9) exceeds the volume of the region.
All subsequent root growth apart from that entering the
granular fertilizer-enriched zones enters unfertilized soil.
When fertilizer P granules are incorporated into soil,
water vapour moves towards them and they dissolve. The
fertilizer P then moves outwards at a rate dependent on soil
moisture content. On the basis of work by Lawton and
Vomocil (1954); Lehr et al., (1959), Burns et al., (1963) and
de Sousa and Volkweiss (1987), it was deduced that
Rgrn ˆ minf3:33Rgrno ‡ 7
0; 0
1f3
33Rgrno
5†
‡ 13
3‰Vl Vl ÿ 0
166†Tg Š0:5 gg
where Rgrn is the radius (mm) of the P-enriched zone
around the granule, and Rgrno is the radius of the fertilizer
granule (mm). Vl is the volumetric water content, and Tg is
time (in days) from incorporating the granule in the soil.
The extent to which this equation depends on soil type is
uncertain.
In studies into the e€ects of fertilizer granule size on crop
response to P, an equation has been derived, by two
independent procedures, for the probable number of
granules intercepted by roots (Moreno, 1959; Costigan,
1987c). Earlier work (Nelder, 1964) had established
equations relating the surface area/volume ratio of an
object to the mean length of chords thrown at random
through it. From these relationships it was derived that the
fraction of the total root length that is within the P-enriched
zones around the fertilizer granules is proportional to the
ratio of the volume of these zones to the total volume of
soil, i.e. to a depth of 30 cm. It was assumed that this
fraction was equal to the fraction of roots absorbing P from
the P-enriched zones. The proportionality constant was
considered to be one, as experimental evidence indicates
that additional root proliferation in soil enriched with P is
generally small unless there is acute P de®ciency in the
rooting medium (e.g. Castilhos and Anghinoni, 1983;
Brouder and Cassman, 1994). The chemical changes within
the P-enriched zones are as described in the bulk soil; the
e€ects of very low pH values at the centre of granules are
ignored.
Plant P uptake. P uptake by roots is determined by ®rst
calculating the maximum amount of P that can be
transported to the roots; this amount is then modi®ed
depending on the plant P concentration (Pgrth) to give P
uptake.
Transport of P is assumed to be solely by di€usion as the
contribution of mass ¯ow is generally negligible (Jungk and
Claassen, 1997). Each day a segment of root is formed in
one or two regions of soil. Also for each day, from their
formation to the end of the growing season, the maximum
amount of P that can di€use to each of them is calculated
284 Greenwood et al.ÐDescription of Model for Crop Response to Phosphate
separately. The calculations are similar irrespective of where
the root segments were formed, but for the sake of clarity,
subsequent descriptions will be for segments in unfertilized
regions of soil. It is considered that the potential ability of
roots to absorb P (e.g. Clarkson et al., 1978) and their
radius do not vary along the length of the root. The
concentration of solution P at the root surfaces is assumed
to be zero as they can absorb P from very low concentra-
tions compared with those in soil solution. In practice, it is
convenient to work in terms of extractable soil P, but this is
converted to solution P through the di€usion coecient as
described later [eqn (12)]. We visualize that each newly
formed root segment is a cylinder, of radius a, with its
surface maintained at zero extractable soil P and that this
cylinder is placed in uniform soil of in®nite extent.
According to this model, it may be derived from Fick's
law (Crank, 1957) that the ¯ux of P to the root surface, Fp is
given by:
Fp ˆ Ds Pextrvo f pGtpy † ÿ0
5 ‡ 0
5 ÿ 0
25 Gtpy =p†0
5
‡ 0
125Gtpy g=a for Gtpy 4 3 6†
Fp ˆ 2Ds Pextrvo f‰ln 4Gtpy † ÿ 2gŠ ÿ1 ÿ g=
‰ln 4Gtpy † ÿ 2gŠ2 g=a for Gtpy 4 3 7†
where Ds is the di€usion coecient through soil [calculated
by eqn (12)], Gtpy a temporary dimensionless coecient
de®ned by the expression Gtpy ˆ Ds Tdf =a2 in which Tdf is
time from the formation of the increment in root length and
g ˆ 0
5772 is Euler's constant. For a considerable period,
Pextrvo is the concentration of extractable soil P (in mg cm ÿ3)
in regions of soil una€ected by root uptake at the time of
formation of the root segment. The value of Pextrvo,
however, is time-dependent because of interchange between
extractable and non-extractable forms of soil P [eqn (11)].
During this period, the P depletion zones of soil, calculated
as will be shown in eqn (9), are regarded as independent of
one another. Eventually, however, if growth continues for a
long time, the total volume of the di€usion zones may
occupy the entire volume of soil available for rooting.
When this happens, Pextrvo, for any newly formed segment is
taken to be the mean concentration of the entire soil
volume.
There is a restriction on uptake by root segments in
regions enriched by granular and starter fertilizers. When-
ever uptake by a root segment exceeds 20 % of the
extractable P in the depletion zone around it, then that
segment ceases to absorb P. This limit was estimated from
reviews of the literature (including De Wit, 1953; Cooke,
1967, p. 262; Wild, 1988, p. 734) which indicated that
arable crops invariably recover no more than 10 % of the
applied fertilizer P in the year of application, although
occasionally recoveries of up to 25 % have been reported.
According to eqns (6) and (7), the ¯ux ®rst declines very
sharply and then slowly with time, resulting in simulations
being sensitive to the integration step. To permit accurate
integration it is necessary to have a means of calculating an
average value of the terms within the brackets (B) over any
time interval that is chosen for integration. This was
achieved by integrating B against Gtpy (using time steps of
0
01 d), ®tting an empirical equation to it, ®nding the area
under the curve corresponding to the values of Gtpy at the
beginning and end of the time step and dividing it by the
di€erence in the two values. With this modi®cation, ¯uxes
were calculated by eqns (6) and (7) with the value of Ds
calculated as the mean value over the life of the segment.
The sum of the ¯uxes to each of the segments is the
maximum possible uptake. The actual amount is less than
this because plant P concentration, at least above a certain
critical level, can suppress P uptake by roots (e.g. Drew et al,
1984; Dunlop et al., 1997; Liu et al., 1998). If Gf is the ratio
of the actual increment in P uptake to the maximum
possible increment then
Gf d† ˆ min 1; e ÿ2
7x † 8†
where
x ˆ Pgrth d†=Popt W†† ÿ Gp
Popt W† ˆ Popt 0† 1 ÿ Kcn W†
where Pgrth(d) is the % P in the plant dry matter and
Popt(W) and Gp are coecients de®ning the suppression of
uptake by % P. Popt(0) is the value of the parameter when
W ˆ 0 and Kcn corrects for its decline with increase in W.
The equation is that of Siddiqui and Glass (1982) modi®ed
to ensure that when the % P in the plant is below a
threshold level, P uptake is una€ected by the % P in the
plant; it is then limited by the amount of P that can di€use
to the roots.
P depletion zone. The daily increase in the volume of the
P depletion zone (DVdpl ) around each daily increment in
root segment, DLdpl(cm), is calculated by
DVdpl ˆ 4pDLdpl Ds d†Tdf 9†
where Ds cm2 d ÿ1 † is the di€usion coecient through the
soil, Tdf is the time in days from the formation of the
increment and DVdpl is in cm3. The equation was derived by
considering that the spread of the di€usion zone normal to
the root's surface is given by (4Dst)0
5 (Tinker and Nye,
2000, p. 133). This relationship may be deduced by
assuming that under uniform conditions the radius of the
depletion zone is equal to the root mean square displace-
ment of a molecule, di€using in two dimensions, from a
point source.
Volumetric soil water content. The average volumetric
soil water content Vl is updated for each day of the
simulation. It is calculated from soil moisture de®cit, the
volumetric water content at ®eld capacity and the depth of
soil from which water is lost by evapotranspiration.
Greenwood et al. (1996) describe the algorithm for soil
moisture de®cit. It requires the daily rainfall, including
irrigation, evaporation from an open water surface and the
current simulated values for soil moisture de®cits and plant
dry weights. Plant dry weights are required both to
determine the fractional crop cover and the depth of soil
 Greenwood et al.ÐDescription of Model for Crop Response to Phosphate
from which water is extracted. Water moves rapidly through
soil so, in contrast to phosphate, only a few roots
penetrating to depth can absorb large quantities of water.
The depth of soil containing 90 % of the roots in cm
(Droot(d)) of non-leguminous crops grown in a sandy loam
soil has been well described by:
Droot d† ˆ 17
6 ‡ 8
53W d†
where W is expressed in t ha ÿ1 of plant dry matter
(Greenwood et al., 1982). It was assumed in the model that
water was extracted to a depth of Droot d† ‡ 10 cm and that
on any day, the volumetric water content was the same
throughout this depth.
Interchange between di€erent forms of P. Equilibrium
between solution P and extractable P is assumed to be
instantaneous. However most of the soil P is in other forms,
including minerals which we group together as non-
extractable P. There is interchange between extractable
and non-extractable P but it is slow and it is re-calculated
for each day for each region and for each zone separately
by:
d Pextrv †=dt ˆ ‰Knextr Pnextrw ÿ Kextr Pextrw ŠBd
where Pextrw and Pnextrw are the extractable and non-
extractable concentrations of soil P expressed on a per unit
weight basis, Knextr and Kextr are rate coecients and Bd is
the bulk density of soil. The equation is a simpli®cation of
Barrow's (1983) solid state di€usion model for sorption and
desorption of phosphate in soil minerals. Experimental
evidence in Fig. 2, discussed later, provides further support
for the validity of the equation.
Di€usion coecient. The di€usion coecient, Ds(d), of
extractable P through soil to the root surfaces is calculated
for each day by:
Ds d† ˆ Dl d†Vl d†Ftort d†dPl d†=dPextrv d†
(Tinker and Nye, 2000) where Pl(d) is the concentration of P
in soil solution, Pextrv(d) is the extractable-soil P and Dl(d) is
the di€usion coecient of P through water after adjustment
for temperature Ttemp. At 25 C; Dl ˆ Dw ˆ 1:02cm2 d ÿ1.
+20
Extractable P
0 100 200
Change in extractable P
0 R(d) ˆ 1. The starting weight was the dry weight of seed
–20
–40
–60
F I G . 2. Relationship between change in extractable P (mg g ÿ1) over
351 d and the level of extractable P at the end of this period.
285
Dl is directly proportional to absolute temperature and
inversely proportional to viscosity. Viscosity, Z in cP, is
approximately related to temperature in degrees centigrade
by
Z ˆ 0
000602 Ttemp †2 ÿ 0
049485 Ttemp † ‡ 1
742 13†
10† which gives a good ®t to the data in Weast (1985). It is
assumed that soil temperature is the same as air tempera-
ture. Ftort(d) is a tortuosity factor for P transport through
soil calculated from Vl(d) the volumetric soil water content,
from
Ftort d† ˆ 1
61 Vl d† ÿ 0
1† for Vl d† 4 0
1
14†
Ftort d† ˆ 0
0001 for Vl d† 4 0
1
which has ®tted measurements on sands and loams
(Barraclough and Tinker, 1981).
The bu€er power, the reciprocal of which is required for
the calculation of the di€usion coecient, is always
assumed to be that in the unfertilized regions of soil and
is calculated by
Gbfr ÿ1†=Gbfr
11† dPextrv =dPl ˆ Gafr Gbfr Pextrv =Gafr † 15†
where Pextrv is the concentration of extractable soil P,
expressed per unit volume of soil, Pl is the concentration of
P in soil solution, and Gafr and Gbfr are coecients
characteristic of the soil. Equation (15) is the derivative of
the Freundlich isotherm:
Pextrv ˆ Gafr PG
l
bfr
16†
which has given good representations of phosphate
adsorption in a range of world soils (Tinker and Nye,
2000). Hysteresis between adsorption and desorption is
ignored.
12†
Parameterization and inputs
Table 1 gives the default values for a typical non-
leguminous vegetable crop grown on a coarse sandy loam.
It will be shown in a companion paper (Greenwood et al.,
2001) that the plant parameters correspond closely with
those of carrot, Daucus carota L.
Plant parameters. Gwt was calculated by solving eqn (1)
300 over the growth period with inputs of daily temperature and
sown (W0) and the maximum value of W at harvest (Wmax).
The secant algorithm for solving non-linear equations
(Johnson and Riess, 1977) was adopted to solve the
equation and always rapidly provided an accurate solution.
Estimation of Phlf is not straightforward. Although many
workers have related growth after a given period to % P,
growth in these measurements is the integral of the e€ects
over time of % P on growth rate, both of which vary;
therefore, these measurements do not provide a satisfactory
basis for estimating Phlf. What is required is the relationship
between relative growth rate (RGR) and % P when both
286 Greenwood et al.ÐDescription of Model for Crop Response to Phosphate
variables are measured at the same time. Both the nutrient
interruption technique (Burns et al., 1997) and nutrient
addition procedures (Ericsson and Ingestad, 1988) appear
to meet these requirements. Burns et al., (1997) grew lettuce
and Ericsson and Ingestad (1988) grew birch seedlings, yet
the relationships between RGR as a fraction of the
maximum against % P are approximately the same. Values
of Pmin(0) ˆ 0
1 % and Phlf (0) ˆ 0
04 % were estimated
from these data. The value of Pmin(0) can be deduced from
many other experiments. For a wide range of crops, Pmin (0)
is generally about 0
1 % (Chapman, 1966; Brewster et al.,
1975; Mengel and Kirkby, 1978; Yost and Fox, 1979;
Stribley et al., 1980; Tinker et al., 1982; Fohse et al., 1988;
Marschner, 1995; Bollons and Barraclough, 1999). Pmax (0)
was estimated as 0
6 % from measurements on lettuce
seedlings grown in fertilizer experiments conducted in the
UK (Costigan, 1986). Kcn is an average value estimated
from measurements of critical % P during growth of a
range of vegetable crops (Greenwood et al., 1980). The root
radius, a ˆ 0
008 cm, is the average of many crops given in
de Willigen and van Noordwijk (1987). The initial % P in
the seed, Pseed , is 0
8 %Ðit is the average of measurements
on seeds for a range of vegetable crops (Haworth and
Cleaver, 1964).
Soil parameters. All soil parameters were determined on
a coarse sandy loam soil of the Wick series at Horticultural
Research International, formerly the National Vegetable
Research Station, Wellesbourne, UK. The soil is described
by Whit®eld (1974).
Values of the rate coecients Kextr and Knextr were
calculated from measurements made by Stone (1998). Eight
di€erent levels of triple superphosphate [Ca(H2PO4)2]
supplying between 0 and 3500 kg P ha ÿ1 were incorporated
in soil 2
5 years before measurements began. Levels of
extractable soil P were measured then, and again 351 d
later. The change in extractable soil P plotted against the
level of extractable soil P for each treatment is shown in
Fig. 2. The relationship is approximately linear which
supports the validity of eqn (11). Substitution of the
gradient of the relationship, divided by 351, in eqn (11)
gives a value for Kextr of 6
37  10 ÿ4 d ÿ1. According to
Fig. 2, d(Pextrw)/dt ˆ 0 and thus equilibrium is reached with
Pextr ˆ 62
3 mg g ÿ1. It follows from eqn (11) that at this
point, Knextr ˆ KextrPextrw/Pnextrw. Substitution in this
relationship of the above values for Kextr and Pextrw and
an estimate of Pnextrw of 1190 mg g ÿ1 gives
Knextr ˆ 3
32  10 ÿ5 d ÿ1.
The coecients, Gafr and Gbfr, de®ning the relationships
between extractable P, Pextrw, and soil solution P, Pl, were
obtained by ®tting the logarithmic form of eqn (16) to
measurements made on samples taken from plots that had
been brought to di€erent levels of Pextrw by incorporation of
a range of fertilizer P levels 3
5 years previously. Pextrw was
determined by extraction with 0
5 M NaHCO3, pH 8
5
(Ministry of Agriculture Fisheries and Food, 1986) and
converted to Pextrv by multiplying by the bulk density, Bd.
The concentration in soil solution, Pl, was determined in
saturation pastes (Hess, 1971). The data are taken from
Table 2 of Stone (1998).
Inputs. To run the model the following inputs are
required:
Weather: Rainfall (including irrigation), evaporation
from an open water surface and mean air temperature for
each day.
Soil: Bulk density, volumetric water content at ®eld
capacity, soil moisture de®cit on some date prior to drilling,
extractable soil P, parameters of an isotherm to de®ne the
relationship between extractable soil P and soil solution P
and the rate constants for interchange between extractable
and non-extractable soil P.
Fertilizer P: The amount of fertilizer P incorporated in
soil, the time of incorporation, the average radius of the
fertilizer granules, amount of starter fertilizer and the
volume of the soil it occupies at the time of injection.
Cultural: Crop species, date of planting and harvest,
initial dry weight (W0) calculated as the product of plant
population and seed dry weight, % P of seed (Pseed) and
potential maximum plant dry weight at ®nal harvest, Wmax.
Table 1 gives default values for all the above inputs
apart from the daily weather which was the best estimate for
Wellesbourne, UK. The method of estimation and the
actual values are given at www.qpais.co.uk/phosmod/
phos.htm
R E S U LT S
Unless otherwise stated, simulations were run with the
default parameter values given in Table 1. Integration was
by Euler's method (Thornley and Johnson, 1990) with time
steps of 1 d because weather inputs were only available for
each day and none of the input values changed rapidly with
time. Running the model with constant temperature and
soil water content provided limited evidence that the
method did not lead to substantial errors; reducing the
step length from 1 to 0
1 d seldom a€ected simulated values
of W or % P by more than 10 %.
Sensitivity analyses
A sensitivity analysis was carried out into the dependence
of plant dry weight, W, on the following parameters: starter
fertilizer, Astfert; granular fertilizer, Agfert; extractable P,
Pextrw; volumetric water content of soil, Vlsens; volume of
soil enriched with starter fertilizer, Vst; the coecients of
the adsorption isotherm, Gafr and Gbfr; the dry weight of
sown seed, W0; the % P of the seed, Pseed; and the rate
constants for the inter-conversion of extractable and
non-extractable soil P, Kextrr and Knextr. These, and the
other inputs to the model given in Table 1, had the default
values given in that table. In addition, temperature is given
a default value of 158C. Each parameter was varied, in turn,
from 0
5±to 2
0±times its default value whilst the other
values were kept constant. The analyses were carried out
with Wmax ˆ 14
5 t ha ÿ1, Th ˆ 146 d; Wmax ˆ 7
0 t ha ÿ1,
Th ˆ 95 d; and Wmax 1
0 t ha ÿ1, Th ˆ 46 d where Th is the
period of growth from the time of planting. The latter two
pairs of values were calculated by ®rst solving the integral
of eqn (1) with Gtemp(d) ˆ 1, R(d) ˆ 1, Wmax ˆ 14
5 t ha ÿ1,
Th ˆ 146 d and W0 ˆ 1
64 kg ha ÿ1 to give Gwt ˆ 0
161 d ÿ1.
 Greenwood et al.ÐDescription of Model for Crop Response to Phosphate 287
6.5 e€ects of increasing Vlsens, Agfert, Pextrrw and Gafr diminished
G afr
with increase in Wmax. The generally small response to Agrfert
over all combinations of other parameters is notable. Vst,
6.0 W0, and Pseed had an appreciable e€ect on response only
Plant dry weight (t ha–1)

when W was very small (1 t ha ÿ1) or when Gafr (and thus the
A gfert adsorption of P) was very high (not shown). Kextr and Knextr
5.5
A stfert only had an appreciable e€ect on W when the extractable P
G bfr was low and there was no starter fertilizer.
5.0 P extrw

4.5 V Comparison between model calculations and published

lsens
experimental results

4.0
0 0.5 1.0
Relative variation in parameter
F I G . 3. Sensitivity of the calculated plant dry weight as each of the
parameters is varied from 0
5- to 2
0-times its default value in Table 1
while the other parameters are held constant. Wmax was set at 7 t ha ÿ1
and Th at 95 d, but all other parameters are as in Table 1.
This value was then used to calculate the dependence of
Wmax on Th throughout growth. Those parameters that had
most e€ect on the responses were identi®ed and varied in
two-dimensional sensitivity analyses.
Overall the most important factors in¯uencing plant dry
weight are Gafr, Gbfr, Pextrw, Vlsens, Astfert and, to a lesser
extent, Agfert, as illustrated in Fig. 3. The main features of
the two-dimensional analyses are given in Table 3. The left-
hand side gives the values of the parameters used in the
simulations and the right-hand side gives the corresponding
e€ect on W of changing the value of the indicated
parameter. Increasing the values of Vlsens and decreasing
Gafr, both of which improved transport of P though soil,
suppressed responses to Astfert and Pextrrw. The bene®t from
Asrfert always diminished with increase in Wmax but responses
to the other parameters changed in a complex manner with
increase in Wmax. However, when Astfert is withheld, the
Long-term experiments have been reported in which
1.5 2.0 some plots were brought to a high level of soil P by
applications of P fertilizer, whilst P fertilizer was withheld
from other plots which thus had a low level of soil P. After a
long period, new treatments consisting of di€erent levels of
fertilizer P were imposed on each set of plots. It was found
that no matter how much fresh fertilizer P was broadcast
and incorporated immediately before drilling, yields were
often lower on the low-P than on the high-P plots (Cooke,
1967; Johnston et al., 1986; Costigan, 1987a). The model
predicts this phenomenon as is illustrated in Fig. 4. The
open symbols in Fig. 4A give the e€ect on yield of
broadcasting and incorporating fertilizer P immediately
before drilling into a low P soil whereas the open symbols in
Fig. 4B give the e€ects of increasing levels of extractable
soil P (which were the residues of fertilizer applications in
previous years). If it is assumed that all the fertilizer P is
extractable then the fertilizer P axis of Fig. 4A is directly
comparable with the extractable P axis in Fig. 4B.
Throughout the range, and even when P levels are
excessively high, fertilizer P treatments gave lower yields
than extractable P treatments. The di€erence is most
marked at the earlier harvest when the maximum yield
obtained with fertilizer P was 4
1 t ha ÿ1 compared with 5
2 t
ha ÿ1 with the extractable P treatments. The model outputs
are therefore consistent with results of ®eld experiments.

T A B L E 3. Summary of two-dimensional sensitivity analyses

Values of parameters used in simulation* Percentage response{ to changes of

Wmax Vlsens Astfert Agfert Pextrw Gafr Vlsens Astfert Agfert Pextrw Gafr

1 0
2 0
01 0
05 18 36
9 9
7 19
4 1
9 10
4 15

7 0
2 0
01 0
05 18 36
9 12
3 12
1 3
2 21
2 33
4
14
5 0
2 0
01 0
05 18 36
9 6
9 6
7 3
8 23
1 31
4
7 0
4 0
01 0
05 18 36
9 ± 3
6 3
1 9
4 11
4
7 0
2 0
01 0
05 36 36
9 5
9 5
8 2
1 ± 14
4
7 0
2 0
01 0
05 9 36
9 34
0 20
5 8
2 ± 45
7
7 0
2 0
01 0
05 18 18
5 5
2 4
3 3
2 11
9 ±
7 0
2 0
01 0
05 18 74
0 33
7 51
8 9
5 52
1 ±
1 0
2 0 0
05 18 36
9 63
9 ± 27
0 111
6 107
4
7 0
2 0 0
05 18 36
9 32
3 ± 15
2 89
8 90
6
14
5 0
2 0 0
05 18 36
9 16
2 ± 9
17 64
7 67
8

*The parameter values in bold type are di€erent from those used in Fig.3
{Calculated, with one exception, as the change in W as a result of increasing the parameter from 0
5- to 2
0-times of its default value and
expressing it as a percentage of the default value. The exception is V1 where the range of values was 1
0- to 2
0- of the default value (virtually no
growth took place with 0
5-times its default value).
288 Greenwood et al.ÐDescription of Model for Crop Response to Phosphate
15.0 15.0
A B

Plant dry weight (t ha–1)

Plant dry weight (t ha–1)
12.5 12.5

10.0 10.0

7.5 7.5

5.0 5.0

2.5 2.5

0.0 0.0
0.00 0.10 0.20 0.30 0.40 0.50 0.08 0.18 0.28 0.38 0.48 0.58

Fertilizer P (t ha–1) Extractable soil P (t ha–1)

F I G . 4. Simulated e€ects on dry matter yield of (A) fertilizer P incorporated into a low P soil immediately before drilling; and (B) extractable soil P
in the absence of added fertilizer. Closed symbols indicate that starter fertilizer (0
01 t P ha ÿ1) is also applied, and open ones that it is not. Values
after 85 d from drilling are represented by triangles and after 146 d by circles. The extractable soil P immediately before fertilizer incorporation is
18 mg g ÿ1 which is equivalent to 0
08 t haÿ1.

12 14 4 5
A B

Plant dry weight (t ha–1)

Plant dry weight (t ha–1)

12
10
Crop response to P

Crop response to P
4
3
10
8
8 3
6 2
6
2
4
4
1
2 1
2

0 0 0 0
0 25 50 75 100 125 150 0 25 50 75 100
Days from drilling Days from drilling

F I G . 5. Dependence on time from sowing of crop response to P (Q) and P-sucient plant dry weight (O). A, Simulated response calculated as
yield with excess extractable soil P (140 mg g ÿ1) divided by the yield when extractable P is low (18 mg g ÿ1). B, Measured response of French beans
calculated as yield with granular fertilizer P of 320 kg ha ÿ1 divided by the yield with no added P. The s.e.m. of the dry weights for the successive
harvests of French beans are 0
0058, 0
0394, 0
169, and 0
205 t ha ÿ1 with 8 d.f.

Experiments have shown that small amounts of starter Fig. 4A) on a low P soil can be as high as those obtained
fertilizer P, positioned in a narrow band so that the roots of with any level of extractable soil P (open symbols, Fig. 4B).
young seedlings intercept it, can dramatically increase Another feature of these two ®gures is that the simulated
yields, even when much granular fertilizer P is incorporated percentage responses to P were much greater after 85d than
in the soil (Costigan, 1987b, c; Costigan and Heaviside, after 146 d. It suggests that the model might predict a
1988; Stone, 1998). Some of the experiments have involved characteristic time course of P response. Consider an
starter fertilizers that contain other nutrients besides experiment in which the default crop was grown on plots
phosphate but it has been shown that the bene®ts generally with high and low levels of soil P . The ratio of growth on
result from the phosphate (Costigan, 1988). Starter the high soil P plots to that on the low soil P plots is a
fertilizers have increased yield dry weights both at the measure of the crop response to excess P. According to the
seedling stage and at commercial maturity even on soils model, once the reserves of seed P have been exhausted, the
with high levels of extractable (0
5 M NaHCO3 pH 8
5) P response to P increases during the exponential period of
(between 40 and 150 mg P kg ÿ1 of dry soil) (Costigan,1988; growth and then decreases during the linear phase, as
Costigan and Heaviside, 1988; Stone, 1998). The model illustrated in Fig. 5A. These features are consistent with
predicts similar bene®ts; starter fertilizer increased yields on anecdotal evidence, but only a few multi-harvest P response
soils containing up to 0
4 t ha ÿ1 extractable soil P which is experiments seem to have been reported in the literature.
equivalent to 90 mg of P g ÿ1 of extractable P (Fig. 4B). It We have therefore included the unpublished results
has also been found in ®eld experiments that starter (Fig. 5B) of an experiment carried out at Wellesbourne,
fertilizer together with broadcast fertilizer P incorporated UK, admittedly on a di€erent crop, French beans
into a low P soil immediately before drilling can increase (Phaseolus vulgaris L.), but they nevertheless give a similar
yields to levels similar to those that can be obtained if the pattern of response to that predicted. In another experiment
soil had high levels of extractable P (Stone, 1998). The with this species, Costigan (1987a) found that response to P
model simulates this e€ect as the simulated dry weights of was considerable 32 d after emergence but had disappeared
plants given starter and granular fertilizer (closed symbols, at commercial maturity. Costigan (1988) and Stone (1998)
 Greenwood et al.ÐDescription of Model for Crop Response to Phosphate
report similar e€ects of time on growth response to starter
fertilizer.
The e€ects of interchange between non-extractable and
extractable soil P [eqn (11)] can be important when
considered over a much longer time scale than in the
sensitivity analyses. If, as is usually the case, the concen-
tration of non-extractable soil P is many times greater than
that of extractable soil P then the values of the rate
coecients, Kextr and Knextr, will remain approximately the
same over a long period. This means that whatever level of
extractable soil P has developed over the years, it will tend
towards a value characteristic of the soil. Thus, if the
extractable soil P had been brought to a high value by
heavy applications of fertilizer P it would subsequently
decline whether or not a crop was grown. Conversely, if the
extractable soil P had been brought to a very low value by,
for example, continuous growth and removal of grass, it
would subsequently tend to rise if the soil was left fallow or
even if some crops were grown on it whilst nutrient
additions were withheld. This phenomenon is illustrated
in Fig. 6, which gives the time course of simulated changes
in extractable soil P and the growth of the default crop in
®ve successive seasons on a soil that initially had a low
extractable soil P content (15 mg P g ÿ1 dry soil). In the ®rst
year growth was poor, but despite uptake of P by the crop
the concentrations of extractable soil P increased through-
out the year. In the second year growth was better and ®nal
yields were almost twice as high as in the ®rst year. Again,
the extractable soil P continued to rise while the soil was
fallow and during the early part of the growing period while
there was little growth, but once dry matter yield had
increased above about 2 t ha ÿ1 (i.e. during the linear phase
of growth) P uptake suppressed extractable soil P. The same
broad pattern was obtained with successive crops. Para-
doxically, both the maximum yields and concentrations of
extractable soil P tended to increase with time despite no
fertilizer having been applied. These are, of course,
predictions from the model, but it is notable that these
trends have been noted in ®eld experiments (Nosko et al.,
1988; Nosko, 1998).
40
30
Extractable soil P
20
10
0 0
0 500 1000 1500
Days from Jan 1st
F I G . 6. Simulated changes in extractable soil P (H) when a crop is
grown in each of ®ve successive years on soil that received no fertilizer
and initially had an extractable P content of 15 mg g ÿ1. The time course
of increase in dry weight of the successive carrot crops is given by (d).
289
DISCUSSION
The rate coecients (Kextr and Knextr) a€ecting the inter-
change between extractable and non-extractable soil P are
the only inputs to the model that are not easy to obtain. In
the sensitivity analyses, however, considerable variations in
their value had little e€ect on P response over the period of
growth of a single crop when extractable soil P was above a
very low value. Thus, by running the model with only very
approximate values for these coecients, it may generally
be possible to get good estimates of P response. It therefore
appears that useful runs of the model can be made with
only readily available inputs.
According to the sensitivity analyses, plant growth was
particularly dependent on starter fertilizer, extractable soil
P, soil water content and the parameters of the adsorption
isotherm. These latter parameters determine the bu€er
capacity which has a large in¯uence on the outputs of other
P models (e.g. Yanai, 1994). The importance of soil water
content is implicit in all di€usion-based models and its
substantial in¯uence on P response in ®eld experiments has
been noted (Greenwood et al., 1974). The considerable
e€ect on P response of raising volumetric water content (Vl )
from 0
2 to 0
4 has not hitherto been emphasized. High
values of Vl will tend to o€set the e€ects of high P
adsorption, such as occurs in peat soils, which may explain
why current UK P fertilizer advice for peat soils is no
di€erent from that for other soils (Ministry of Agriculture
Fisheries and Food, 2000).
The superiority of extractable soil P, Pextrw, and starter
fertilizer, Astfert, over granular fertilizer, Agrfert, in increasing
yields, as revealed by the sensitivity analyses, warrants
discussion. Such analyses can only cover a small fraction of
the possible conditions so conclusions may be atypical.
However, this possibility is unlikely as the simulated
responses were similar to those found in past experiments
(Fig. 4). To explore the reasons for the di€erences in
e€ectiveness of these di€erent forms of P, ®rst consider the
time course of growth and crop P uptake. When growth is
not limited by P, the model calculates that dry weight ®rst
increases almost exponentially and then linearly with time.
In consequence, the rate of dry matter increase is initially
20
almost proportional to the plant dry weight and then almost
constant. The plant P concentration needed for optimal
Plant dry weight (t ha–1)
15 plant growth, however, decreases with increase in plant
weight. The net e€ect is that the rate of P uptake needed to
sustain maximum growth ®rst increases sharply with time,
10 reaches a maximum, and then declines slowly. Meanwhile,
the total length of root that is capable of absorbing P from
the soil increases throughout the growing period. Thus, as
5
has been suggested by Scaife (1994), the ability of roots to
supply P relative to crop demand will increase during the
linear phase of growth. Crop response should rise to a
2000
maximum and then decline as the plant grows, as is
predicted by the model and as is found experimentally
(Fig. 5). Crops will be most vulnerable to P de®ciency in the
early stages of growth. A temporary set-back at that time
could restrict ®nal yield even if there was ample P in the
plant for the remainder of the growing period. Only if the
290 Greenwood et al.ÐDescription of Model for Crop Response to Phosphate
plant could compensate, for example by delaying commer-
cial maturity, would ®nal yields be una€ected.
Starter fertilizer, by relieving P de®ciency in the very early
stages of growth and thus the set-back that often occurs,
increases ®nal yield considerably. Broadcasting and incor-
porating granular fertilizer immediately before drilling is not
e€ective in relieving P de®ciency in the early stages because
such a small proportion of the total root length intercepts P-
enriched zones and is thus able to absorb P. Raising the
overall level of extractable soil P is better because it improves
P supply to the entire root system. The indication that the
extractable P may tend toward an equilibrium level
characteristic of the soil [eqn (11) and Fig. 6]) suggests
that if more P fertilizer is applied than is needed to maintain
it at this level, much of it will be wasted. It will be converted,
at a rate depending on the soil, into non-extractable soil P
which is largely unavailable to subsequent crops. Ideally, it
would be best to maintain the soil at the equilibrium level by
applications of manures and fertilizers. This practice would
not permit high yields on some soils, but on others, such as
that described in this paper, it could do so. For soils of this
latter type, an ecient way of ensuring adequate crop
nutrition may be to maintain the extractable soil P at the
equilibrium level for the soil, and then to inject starter
fertilizer for demanding crops. To further develop this
theme, there is a need to devise a more rapid means of
determining the relationship between extractable and
solution P and the rate coecients between extractable
and non-extractable soil P. In addition, there is a need to
calibrate crop parameters for di€erent species; this is the
topic of the following paper (Greenwood et al. 2001).
AC K N OW L E D G E M E N T S
The authors thank Dr Marcel van Oijen for helpful comments
on an earlier draft of the paper.
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