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13 Comparison of Pig Growth Models

– The Genetic Point of View
P. LUITING AND P.W. KNAP
PIC International Group, Ratsteich 31, D-24837 Schleswig, Germany
pieter.knap@pic.com

Introduction

Application models of growth in pigs were pioneered by Whittemore and

Fawcett (1976) and further developed by Moughan and Smith (1984),
Black et al. (1986), Emmans (1988, 1997), Pomar et al. (1991), TMV
(1991), De Greef (1992), Ferguson et al. (1994), Whittemore (1995), and
many others. All these models are dynamic (describing the growth
process over time), deterministic (non-stochastic), and semi-mechanistic
(more or less based on biological growth mechanisms, focusing on the
accretion of several body components). Many of these biological
mechanisms are driven by one or more parameters that are specific to
the genotype (breed, strain, etc.) to be simulated. Proper simulation of
that genotype then requires quantification of those parameters, which
requires experimental effort. Therefore, these parameters are of primary
interest for animal breeders who use the model for their particular
purposes.
The above mentioned publications describe the mechanisms that
form the core of their models with different notation and
argumentation; as a consequence, the exact differences and similarities
between these approaches are often unclear and we attempt here to
clarify some of these issues. Section 1 of the present text describes the
main common mechanisms of the various models and their genotype-
specific aspects. Section 2 describes the details of implementation of
these mechanisms, and attempts to derive a common denominator.
Section 3 discusses the differences between the various models from the
point of view of adequacy and complexity of the description of the
genotype.
Given the large amount of water in the mammalian body, the
prediction of body water mass is perhaps the most critical factor of pig

© CAB International 2006. Mechanistic Modelling in Pig and Poultry Production

(eds. M. Freer and H. Dove) 260
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Comparison of Pig Growth Models 261

growth simulation. Emmans and Kyriazakis (1995) focus on this issue in

quantitative detail, and this is the only such treatment of the matter that we
are aware of; we do not deal with it any further.

Main Common Mechanisms

All the models discussed here distinguish three processes that require
energy: protein deposition (PD), body maintenance, and lipid deposition
(LD), all three on a daily basis. The driving factor is PD; it is predicted
first, after which it determines all other factors in the simulation. The
metabolizable energy (ME) requirements for body maintenance (MEm) are
predicted from metabolic body weight or from body protein mass: MEm =
α × BW or MEm = α × P, where α and  are constants. It follows that at
known energy intake (MEI), LD is predicted as the item that balances the
requirements of the law of conservation of energy:

MEI = ME + PD × E / k + LD × E / k ?LD =
m P P L L
MEI - ME − (PD × E / k ) (13.1)
m P P
E / k
L L
where LD = lipid deposition (kg per day), MEI = ME intake (MJ per day),
MEm = ME required for maintenance (MJ per day), PD = protein
deposition (kg per day), EP and EL = combustion energy contents of body
protein and lipid, respectively (MJ per kg), and kP and kL = efficiencies of
use of ME for PD and LD, respectively.
All models assume EP, EL, kP, kL, α and  to be constant, although they
vary somewhat with regard to their values. Genetic variation in these
parameters is presumed absent.
All models calculate water and ash deposition from PD, using empirical
equations with constant parameter values. Empty body weight gain (BWG)
is calculated by summation of the four components; adjustment for a
certain amount of gut fill leads to daily growth:

empty BWG = PD + LD + water deposition + ashdeposition? (13.2)

BWG = (empty BWG) × (1 + gutfill correction factor)
All models define an initial status with the starting values for body
protein, lipid, ash and water mass. The predicted deposition values are
added to the current values of body component mass, and next day’s
calculation is performed. This process is repeated until the desired end
status is reached. The models vary somewhat with regard to the water-
and ash-related empirical equations and/or their parameter values, with
regard to the magnitude of gut fill, and with regard to the initial body
component mass assumptions. Genetic variation is presumed absent
here.
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262 P. Luiting and P.W.Knap

Most models treat feed intake as a known input parameter. However,

some models predict voluntary feed intake as an output parameter based
on MEm, desired PD, desired LD and equation 13.1. The prediction of
desired PD and desired LD is described, respectively, in the two sections
below.

Prediction of Protein Deposition

The main feature of PD is that it is sourced from the digestible crude

protein intake (DCPI). Like MEm, protein requirements for body
maintenance (Pm) are predicted from metabolic body weight or from body
protein mass: Pm = γ × BWδ or Pm = γ × Pδ, where γ and δ are constants. It
follows that at known DCPI, PD can be predicted as:
PD = eP × (DCPI − Pm ) (13.3)
where eP = material efficiency with which protein is utilized above
maintenance, DCPI = digestible crude protein intake (kg per day), and Pm
= protein required for body maintenance (kg per day).
The material efficiency with which protein can be utilized above
maintenance (eP) is assumed to have a constant maximum value (maxep) of
about 0.83, see Fig. 13.1. The various models vary somewhat with regard
to the magnitude of this maximum value for eP, and of γ and δ. Genetic
variation is again presumed absent.
All models described here recognize two situations where eP is lower
than its maximum value, i.e. where part of the protein supply is used as a
source of energy rather than amino acids: (i) PD is at its maximum (section
1) and (ii) ME supply is insufficient (section 2).

PD

ep = maxep
Pm DCPI

Fig. 13.1. Protein deposition (PD) in relation to digestible crude protein intake (DCPI). Pm
denotes the protein requirement for body maintenance, eP is the material efficiency of protein
utilization. In this range of DCPI, eP attains its maximum value maxeP.
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Comparison of Pig Growth Models 263

Maximum protein deposition

All models assume a genetically determined upper limit to a pig’s rate of

protein deposition: maxPD. This is the protein deposition that the pig
seeks to realize: maxPD equals the pig’s desired level of PD. By analogy,
the DCPI level that is required to just achieve maxPD is the desired DCPI:
output PD maxPD
eP = = = = max eP
input DCPI − Pm desired DCPI − Pm

When realized DCPI exceeds desired DCPI, the surplus of digested

protein is used as an energy source. This reduces the material efficiency eP,
calculated as:
output PD maxPD
eP = = = < max eP , for DCPI > desired DCPI (13.4)
input DCPI − Pm DCPI − Pm
Adding this relation to Fig. 13.1 results in Fig. 13.2. The various models
use different definitions for the genetic factor(s) that describe maxPD.
The obtained value for maxPD is used to determine desired DCPI =
maxPD
+ Pm, which is compared to DCPI. If DCPI < desired DCPI, =
max eP
PD is estimated with equation 13.3; otherwise, PD = maxPD.

Insufficient energy supply

All models assume implicitly an intrinsic lower limit to a pig’s ME

requirements in order to reach maxPD: minMEImaxPD. From the law of
conservation of energy, all models then assume an intrinsic lower limit to a
pig’s LD coinciding with maxPD:

PD

maxPD

ep < maxep

ep = maxep
Pm DCPI

Fig. 13.2. An extension of Fig. 13.1. PD has an upper limit (maxPD), reached at DCPI =
desired DCPI. At higher DCPI levels, eP is lower than its maximum value.
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264 P. Luiting and P.W.Knap

minMEImaxPD = MEm + (maxPD × EP / kP ) + (minLDmaxPD × EL / kL ) (13.5)

where minLDmaxPD is the lower limit to LD coinciding with maxPD.
The relation of PD and LD with MEI is shown in Fig. 13.3 . At MEI ˇ”δ
minMEImaxPD, LD must compete for resources with PD so its realized level
is an indication of the genotype’s priorities for this resource allocation. At
MEI > minMEImaxPD all surplus energy goes to LD, and this is genotype-
independent. So we have a genetically controlled part for LD below
minMEImaxPD and a genetically controlled part for PD above it. It follows
that the point of intersection represents (i) the pig’s ambition for protein
deposition and (ii) its genetic priorities for resource allocation between PD
and LD, and minMEImaxPD must then be the lower limit to the energy
intake that the pig seeks to realize.
Another consequence is that PD is not the only genetically determined
factor that drives the system: below minMEImaxPD, the desired LD level is
just as important. Thus a genotype must be characterized by a desired PD
(= maxPD) and a desired LD coinciding with it (= minLDmaxPD).
Dependent on environmental conditions, it may well be that neither of
them ever gets phenotypically expressed.
Whereas at MEI > minMEImaxPD the surplus energy is fully used for
LD, the energy deficit at MEI ˇ”δ minMEImaxPD is not fully accounted for
by reducing LD but also by reducing PD. Part of DCPI is then used as an
energy source and not for protein deposition. This leads to PD < maxPD,
and it reduces eP to:
output PD PD
eP = = = < max e for PD < maxPD (13.6)
input DCPI − Pm desired DCPI − Pm P

Adding this relation to Fig. 13.2 results in Fig. 13.4.

PD LD

maxPD

LD

minLD maxPD
PD

b1

b0 minMEI maxPD MEI

Fig. 13.3. Protein (PD) and lipid deposition (LD) in relation to ME intake (MEI). For PD <
maxPD, PD = f(MEI) = b1 × (MEI – b0). Above that level (i.e. above MEI = minMEImaxPD and
LD = minLDmaxPD), all surplus energy is used for LD.
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Comparison of Pig Growth Models 265

PD

maxPD

PD =
f(MEI)
ep < maxep

ep < maxep
ep = maxep
Pm DCPI

Fig. 13.4. An extension of Fig. 13.2. At insufficient ME supply, PD has an upper limit that
depends on ME intake (MEI) and eP is lower than its maximum value.

The various models use different definitions of the function f(MEI):

PD = b1 × (MEI – b0) that describes the reduction of PD at MEI <
minMEImaxPD, see the section on ‘Insufficient energy supply’ in ‘Model
comparison’ above.
The value for PD that was obtained from DCPI (in the section on
‘Maximum protein deposition’ in ‘Main common mechanisms’ above)
PD (from DCPI)
is used to determine desired MEI = + b0, which is
b1
compared to MEI. If MEI < desired MEI, PD is estimated via f(MEI);
otherwise, PD from DCPI remains unchanged.

Model Comparison

The main issue from the above section is: body protein deposition (PD)
takes place at a low material efficiency when it is limited either by the
genetic maximum to PD or by insufficient energy supply. The question is
then how the various models parameterize these relations. We deal next
with maximum protein deposition, with insufficient energy supply, and
give a brief comparison of how some of these models describe voluntary
food intake.

Maximum protein deposition

Three different methods are used for the definition of maxPD.

The simplest method makes the assumption that maxPD has a constant
value throughout the growth period, at least up to commercial slaughter
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266 P. Luiting and P.W.Knap

weights (Whittemore and Fawcett, 1976). Hence, different genotypes can

be characterized by different values for this single parameter. The other
two methods make maxPD depend on body weight or stage of body
development, describing a pattern where maxPD increases rapidly early in
life, plateaus during the growth period and then decreases towards zero at
maturity. Two types of functions are used to model this trend, as shown in
Fig. 13.5.
First, hyperbolic or parabolic equations of maxPD in terms of BW, in
both cases with two genetic parameters.
Walker and Young (1993) used a skew-hyperbolic relation with BW:

maxPD = BW  e(c + d  BW) (13.7)

Thorbek (1975) and Whittemore and Fawcett (1976) used a quadratic

relation with metabolic body weight:

maxPD = f  MBW2 – g  MBW (13.8)

Second, derivatives of sigmoid curves for potential protein growth, based

upon the idea that the effect of BW on maxPD reflects the effect of
physiological development. Emmans (1988) used the Gompertz function;
this equation has two genetic parameters, the rate parameter BGomp and
the mature body protein mass Pmature:
P  P 
maxPD =BGomp × P × ln(Pmature / P) =BGomp × Pmature × × ln1 /  (13.9)
Pmature  Pmature 
Black (1988) used the Richards function, modified to replace one of the
two instances of protein mass by BW:
 P 
maxPD = k × (BW+S)a × [(Pmature P) / Pmature ]= k × (BW + S)a × 1 −  (13.10)
 Pmature 

This equation has a high information requirement for characterization of

the genotype: four genetic parameters (k, S, a and Pmature).

maxPD maxPD
Thorbek (1975)
eqn (8)

Emmans Black (1988)

Walker & Young (1993) eqn (9) eqn (10)
eqn (7)
BW P/Pmat

Fig. 13.5. Various functions to describe the course of maxPD in relation to body weight (BW)
or body protein mass (P) as a proportion of its mature value (Pmat).
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Comparison of Pig Growth Models 267

Insufficient energy supply

All models described here use a function f(MEI) that describes the
reduction of PD for MEI < minMEImaxPD. This function is not always
explicit, but may follow implicitly from other, explicitly defined, relations.
Table 13.1 introduces these structures. Method 1 explicitly defines a value
for minLD/PD, and implicitly describes f(MEI) as a linear regression line
of PD on MEI with b0 = MEm. Methods 2 and 3 define f(MEI) explicitly
but differently. Methods 4 and 5 explicitly define a function for eP and a
value for minL/P, respectively, leading implicitly to a function PD =
f(MEI).
Method 1 was published earliest, and some subsequent methods give,
in addition to their explicit element f(MEI), also their own implicit value
for minLD/PD, for comparison purposes. We include minLD/PD in Table
13.1 for the same reason, although it is not required for a comparison of
these five methods when f(MEI) has already been described.
The general form of f(MEI) is PD = b1 × (MEI – b0), where b0 is an x-
intercept and b1 represents the reduction of PD at MEI < minMEImaxPD.
The general form of minLD/PD is derived as follows. Substituting f(MEI)
into equation 13.1 gives:

MEI − ME − (PD × E / k ) MEI − ME − [b × (MEI − b ) × E / k ]

LD = m P P = m 1 0 P P =
E / k E / k
L L L L

1− b ×E / k  ME − b × b × E / k 
= 1 P P ×  MEI − m 0 1 P P
E / k  1− b ×E /k 
L L  1 P P 

so that

Table 13.1. Methods for the reduction of protein deposition at MEI < minMEImaxPD.
Method minLD/PD PD = f(MEI) Third function
Whittemore explicit definition implicit description
1
Moughan constant → Lin(MEI)
Black implicit description explicit definition
2
De Greef nonLin(MEI) ← Lin(MEI)
3 Van Milgen implicit description ← explicit definition
nonLin(MEI) Quad(MEI)
Emmans & implicit description ← implicit description ← explicit definition
4
Kyriazakis nonLin(MEI, DCPI) Lin (MEI), nonLin(DCPI) eP = Lin(MEI/DCPI)
5 De Lange implicit description ← implicit description ← explicit definition
nonLin(MEI) Lin(MEI, L, P) minL/P = constant
Lin(): linear function. nonLin(): nonlinear function. Quad(): quadratic function.
References in the text.
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268 P. Luiting and P.W.Knap

1 − b1 × EP / kP  MEm − b 0 × b1 × EP / kP 
×  MEI − 
EL / kL  1 − b1 × EP / kP 
minLD / PD = =
b1 × (MEI - b0 )
(13.11a)
MEm − b 0 × b1 × EP / kP
MEI −
1 − b1 × EP / kP 1 − b1 × EP / kP
= ×
b1 × EL / kL MEI − b0

which can usefully be rearranged as

1 − b1 × EP / kP  MEm − b 0 1 
minLD / PD = × 1 − ×  (13.11b)
b1 × EL / kL  1 − b1 × EP / kP MEI − b0 

Method 2 introduced the parameter ‘marginal minLD/PD’ (margminLD/PD):

the ratio between the regression coefficients of LD and PD on MEI, at MEI <
minMEImaxPD. We show in Appendix 13.1 that this parameter equals the first
1 − b1 × EP / kP
term in equation 13.11b: margminLD/PD = . We present
b1 × EL / kL
the implicit values of margminLD/PD as well as minLD/PD for the purpose of
comparison between the models.
The methods are discussed in detail in the following five sections and
in Table 13.2.

Table 13.2a. The forms of regression coefficientsa b0 and b1 for the methods in Table 13.1.
The forms of margminLD/PD and minLD/PD are in Table 13.2b.
Method b0 b1
1 Whittemore MEm 1
Moughan EP / kP + minLD / PD × EL / k L

Black
2 g1 g2
De Greef
g2 × (MEI − MEm − g1) =
3 Van Milgen MEm −4 × maxPD
= × (MEI − MEm − g1)
g12

Emmans & DCPI − Pm

4 0 ×
Kyriazakis DCPI

1
5 De Lange MEm + (minL/P × P – L) × EL / kL
EP / kP + minL / P × EL / k L

aPD = b1 × (MEI – b0).

References in the text.
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31/1/06

Table 13.2b. Continuation of Table 13.2a: the forms of margminLD/PD and minLD/PD for the methods in Table 13.1.
1− b1 × EP / kP  MEm − b0 1 
Method margminLD / PD = minLD / PD = margminLD / PD × 1− × 
b1 × EL / kL
10:20

 1− b1 × EP / kP MEI − b0 

1 = minLD/PD g1

1− g2 × EP / kP  MEm − g1 1 
2 ×
Page 269

margminLD / PD × 1− 
Comparison of Pig Growth Models

g2 × EL / kL  1− g2 × EP / kP MEI − g1 

 
1  1 
3 × − EP / kP  margminLD/PD
EL / kL  −4 × maxPD × MEI − ME − g
( m 1) 
 g12 

 
1 1 DCPI   MEm 1 
4 × × − EP / kP  margminLD / PD × 1− × 
EL / kL   (DCPI − Pm )   1−  × (DCPI − Pm ) × E / k MEI 
P P
 DCPI 

margminLD / PD ×

5 minL/P  (minL / P × P − L) × (EP / kP + minL / P × EL / kL ) 1 

1+ ×
 minL / P MEI − [MEm + (minL / P × P − L) × EL / kL ] 
269
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270 P. Luiting and P.W.Knap

1. Whittemore and Fawcett (1976)

These authors assumed that an intrinsic minimum amount of lipid
deposition must accompany each unit of protein deposition, implying that
there is an intrinsic lower limit to the ratio between lipid and protein
deposition (minLD/PD), which is defined as a genetically determined
constant throughout the growing period (g1 in Table 13.2).
Substitution of minLD/PD × PD for LD into equation 13.1 shows that
f(MEI) is assumed to be a linear regression of PD on MEI:

MEI = MEm + (PD × EP / kP ) + ( minLD / PD × PD × EL / kL )?

1 (13.12)
PD = × (MEI − MEm )
EP / kP + minLD / PD × EL / kL

It follows that b0 = MEm, b1 = 1 / (EP / kP + minLD/PD × EL/kL). The

linear regression coefficient b1 is therefore also assumed to be a genetically
determined constant throughout the growth period.

2. Black et al. (1986)

Black explicitly assumed f(MEI) to be a linear regression of PD on MEI:
PD = b1 × (MEI – b0), and b0 and b1 are assumed to be genetically
determined constants throughout the growth period (g1 and g2 in Table
13.2). In contrast to Whittemore’s model (equation 13.12), the x-intercept
is not assumed to be in MEm; all its estimated values are lower than MEm.
The argumentation for b0 < MEm is that when MEI = MEm, protein
deposition is often positive and lipid deposition is often negative (i.e. lipid
is catabolized). Moreover, Black’s model does not require an intrinsic
minimum amount of lipid deposition that must accompany each unit of
protein deposition. Hence minLD/PD is not a constant; it decreases when
MEI decreases, see Table 13.2. It is genetically determined through b0 and
b1. By contrast, margminLD/PD is a constant in this model.
The entity margminLD/PD (the ratio between the regression
coefficients of LD and PD on MEI) was introduced by De Greef (1992) as a
genetically determined constant throughout the growth period. We show in
Appendix 13.1 that, because of equation 13.1, the regression of LD on MEI
can be fully expressed in terms of the one for PD, which removes the need
for De Greef ’s two additional genetic parameters (e.g. c0 and c1 in LD = c1
× [MEI – c0]). This makes De Greef ’s model collapse entirely to Black’s one;
the only novelty is that it makes the margminLD/PD parameter explicit.

3. Van Milgen et al. (2000)

Like Black, these authors explicitly assumed a regression function for
f(MEI): PD = b1 × (MEI – b0), with an x-intercept b0 = MEm just as in
Whittemore’s model. Their regression coefficient b1 is in itself a linear
regression on (MEI – MEm) and can be written, in our notation, as b1 =
g2 × (MEI – MEm – g1). This way, f(MEI) has the form of a quadratic
regression of PD on (MEI – MEm):
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Comparison of Pig Growth Models 271

PD = b1 × (MEI − b0 ) = g 2 × (MEI - MEm − g1 ) × (MEI − MEm ) =

(13.13)
= g 2 × (MEI − MEm )2 − g1 × g 2 × (MEI − MEm )

Parameters g1 and g2 are defined as genetically determined constants.

Equation 13.13 is defined as having its maximum value at maxPD; this
makes one of the two parameters (g1, g2) redundant, as follows.
Solving equation 13.13 for dPD / d(MEI – MEm) = 0 gives
minMEImaxPD:

dPD
= 2 × g2 × (MEI - MEm ) − g1 × g 2 = 0 (13.14)
d(MEI - MEm )
After substitution of minMEImaxPD from equation 13.14 for MEI in
equation 13.13, maxPD can be solved for in order to express b1 in terms of
maxPD and g1 (instead of g1 and g2):

maxPD = g 2 × (minMEImaxPD − MEm )2 − g1 × g 2 × (minMEImaxPD − MEm ) =

−4 × maxPD
= g 2 × (g1 / 2)2 − g1 × g 2 × (g1 / 2) = −0.25 × g 2 × g12?g 2 = →
g12
dPD
= 2 × g2 × (MEI - MEm ) − g1 × g 2 = 0
d(MEI - MEm )
(13.15)
g1 × g 2
?minMEImaxPD − MEm = = g1 / 2
2 × g2
It follows that b1 is not a constant but a function of MEI. It is genetically
determined through maxPD and g1.
Like b1, minLD/PD is not a constant but a function of MEI; it is
genetically determined through maxPD and g1 (see Table 13.2). Note that
in this model, margminLD/PD equals minLD/PD and is therefore not a
constant either, in contrast to model 2.

4. Kyriazakis and Emmans (1992)

The reduction in eP with decreasing MEI given a certain amount of DCPI
was represented by these authors in terms of a linear regression of eP on
the ratio between the food ME content and food digestible protein content
(MEC / DCPC, which equals MEI / DCPI), forced through the origin:
MEI PD MEI DCPI − Pm
eP =  × → =× → PD =  × × MEI (13.16)
DCPI DCPI − Pm DCPI DCPI

The parameter µ is explicitly assumed to be a constant, not genetically

determined. Following equation 13.16, f(MEI) has the form PD = b1 ×
(MEI – b0) with b0 = 0 and ˇ÷embed Equation.3 ˇ≠ ˇˇ§. The regression
coefficient b1 is then a non-linear function of DCPI, not genetically
determined. Table 13.2 shows that minLD/PD is not a constant, but a non-
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272 P. Luiting and P.W.Knap

linear function of MEI and DCPI, not genetically determined. It also shows
that margminLD/PD is not a constant either, but a non-linear function of
DCPI, not genetically determined.

5. De Lange (1995)
De Lange assumed that an intrinsic minimum amount of lipid mass (L)
must accompany each unit of protein mass (P), hence that there is an
intrinsic minimum lipid to protein ratio in the body (minL/P), which is
genetically determined. Therefore, minLD can be written as minLD =
minL/P × (P + PD) – L. The equivalent of equation 13.12 is then:

PD =
1
EP / kP + minL / P × EL / kL
(
× MEI − [MEm + (minL / P × P − L) × EL / kL ]) (13.17)

1
So b0 =MEm +(minL / P × P – L) × EL / kL, and b1 =
EP / kP + minL / P × EL / kL

It follows that f(MEI) is implicitly defined as a linear regression with a

regression coefficient b1 that is a genetically determined constant, just as
minL/P. Its x-intercept b0 is a function of P and L, and it is genetically
determined through minL/P. When the actual amount of body lipid exceeds
its intrinsic lower limit, then (minL/P × P – L)  0, so that b0  MEm.
Table 13.2 shows that minLD/PD is a function of P and L, and (non-
linearly) of MEI; it is genetically determined through minL/P;
margminLD/PD is a constant, this time equal to minL/P.

Prediction of voluntary food intake

Two of these models predict voluntary food intake, and they do so from
the combination of maintenance requirements and desired PD and LD.
Desired LD is defined in both cases via an additional equation. The forms
of these equations (and their difference between the two models) are
similar to the associated definitions of maxPD, as follows.
Emmans (1988) used again the derivative of the Gompertz growth
function to describe desired LD. This equation has two genetic parameters:
the rate parameter BGomp (equal to the one for maxPD in equation 13.9),
and mature body lipid mass Lmature.
desired LD = BGomp  L  ln(Lmature / L) (13.18)
Black (1988) used again the derivative of a modified Richards growth
function to describe the desired increase of net energy (NE) in the body:
desired NE = k × (BW + S)a × [(NEmature – NE) / NEmature]
which gives
desiredNE − maxPD × EP
desiredLD = (13.19)
EL
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Comparison of Pig Growth Models 273

Discussion and Conclusions

Apart from several empirical relations, the models described here contain
three truly mechanistic elements in the form of equations 13.1, 13.2 and
13.3. All genetic factors enter this mechanistic system through eP in
equation 13.3: maximum protein deposition (maxPD) and protein
deposition at insufficient energy supply.
The differences and similarities between the various models with
regard to these genetic factors are summarized and discussed below.

Maximum protein deposition

The genetic aspects of the five discussed methods are summarized in

Table 13.3.
Black et al. (1986), Moughan and Verstegen (1988) and Emmans and
Kyriazakis (1997) refer to several studies that show an effect of body weight
on maxPD. Although measurement of maxPD seems to be difficult and not
very reliable, especially in the early increasing phase, the trend of
‘increasing rapidly early in life, plateauing during the growing-fattening
stages and then decreasing towards zero at maturity’ is generally accepted.
Hence inclusion of this effect of body weight on maxPD makes the model
valid over a wider growth trajectory than with the assumption of a single
constant maxPD value as in method i in Table 13.3.
The hyperbolic, parabolic and sigmoid-derived equations are all
empirical, and it is probably possible to find functions from all these three
classes that fit the observed pattern very well. However, the derivatives of
the sigmoid functions have a more general character and useful properties
like allometry. Disadvantages of Black’s modified Richards curve (method
v) are the large number of genetic parameters and the unclarity of his
modification. The inclusion of BW has strange consequences; for example,
if a pig has a higher BW at a given P because it has more fat in its body,
then its predicted maxPD will be higher. The derivative of the Gompertz

Table 13.3. Methods to describe maximum protein deposition.

Method na maxPD Type
Whittemore
i 1 g1 constant
Moughan
Whittemore
ii 2 f(g1, g2) hyperbola
Thorbek
iii Walker & Young 2 f(g1, g2) skewed parabola
iv Emmans 2 f(BGomp, Pmature) Gompertz
v Black 4 f(k, S, a, Pmature) modified Richards
aNumber of genetic parameters.
References in the text.
13MechModPig 31/1/06 10:20 Page 274

274 P. Luiting and P.W.Knap

curve with two genetic parameters (method iv) seems to be the preferred
model. An additional advantage is the smooth connection with Taylor’s
genetic size scaling principles, see Emmans (1988, 1997).

Insufficient energy supply

The genetic aspects of the five discussed methods are summarized in Table
13.4 and Fig. 13.6.
De Greef and Verstegen (1992) showed that minLD/PD decreases with
decreasing MEI. This makes method 1 (with a single constant value for the
genetic parameter minLD/PD, g1 in Table 13.4) unsuitable.

Table 13.4. Methods to describe protein deposition at MEI < minMEImaxPD.

Method na b0 b1 margminLD/PD minLD/PD
Whittemore
1 1 MEm g1 f(g1) g1
Moughan
Black
2 2 g1 f(g2) g2 f(g1, g2, MEI)
De Greef
3 Van Milgen 1 MEm f(g1, maxPD, MEI) f(g1, maxPD, MEI) f(g1, maxPD, MEI)
Emmans and
4 0 0 f(DCPI) f(DCPI) f(DCPI, MEI)
Kyriazakis
5 De Lange 1 f(g1, P, L, MEm) f(g1) f(g1) f(g1, MEI)
aNumber of genetic parameters.
References in the text.

PD PD
high L/P

maxPD maxPD

3 4
1
L/P > minL/P

high
DCPI

low
1 DCPI
3
4
2 L/P = minL/P

0 MEm MEI 0 MEm MEI

b0 < MEm b0 < MEm

Fig. 13.6. Methods to describe protein deposition with insufficient ME supply, as in Table 13.3.
Left: (1) Whittemore, Moughan; (2) Black, De Greef; (3) Van Milgen and Noblet; (4) Emmans
and Kyriazakis. Right: De Lange. References and explanation of symbols in the text.
13MechModPig 31/1/06 10:20 Page 275

Comparison of Pig Growth Models 275

Table 13.2 shows that method 2 makes minLD/PD dependent on MEI,

for which it requires two genetic parameters (g1 and g2 in Tables 13.2 and
13.4). The reason given by Black et al. (1986) for this is that when ME
intake is just sufficient to achieve zero energy retention (maintenance
energy intake, MEm), protein deposition is positive and lipid deposition is
negative (i.e. lipid is catabolized). This feature has been emphasized by
many authors (e.g. ARC, 1981; Close and Fowler, 1982; Walker and Young,
1993; NRC, 1996; Van Milgen and Noblet, 1999). Thus, the line describing
PD as a linear function of MEI has an x-intercept lower than MEm (b0 <
MEm): the assumption that the ME requirement at zero PD equals MEm (b0
= MEm) is therefore invalid.
Method 3 incorporates the effect of MEI on minLD/PD in a different
way. It requires two genetic parameters, one of which was shown, in
equation 13.15, to collapse to maxPD; therefore Table 13.1 gives only g1.
This is an advantage over method 2 with its two required genetic
parameters. However, f(MEI) is a quadratic regression of PD on MEI with
an x-intercept equal to MEm (see line (3) in Fig. 13.6), which is (like for
method 1) in conflict with the above mentioned finding of b0 < MEm. This
makes method 3 also unsuitable.
Method 4 does not include any genetic parameters, which makes it an
inconvenient method to model animals that differ in their genetically
desired lipid deposition at MEI < minMEImaxPD. This may be caused by
the fact that this method was based on data collected on very young pigs
(4, 6 or 8 weeks after 12 kg body weight, up to 20–50 kg end weight).
Emmans (1997) notices that ‘all genotypes are thin when they are very
immature’ and that this means that the body lipid to protein ratio shows
little variation at a low degree of maturity. This should make it very
difficult to detect genetic variation in this trait at that stage.
Method 4 presents two more complications. First, when an efficiency
parameter (i.e. output/input) such as eP is expressed in terms of MEI, it
would make sense to regress eP on MEI / (DCPI – Pm) where the
denominator properly represents ‘input’. However, the regression was
done on MEC / DCPC, which equals MEI / DCPI. Of course, Pm is small in
comparison to DCPI (especially in very young animals) so the error is
probably small.
Second, the above regression was forced through the origin, defining
b0 = 0. This approach has one parameter less to be estimated, an
important advantage when analysing a small dataset. However, such an
assumption about an intercept (with notoriously large standard errors of
the regression estimates) is very difficult to falsify, and the default
approach should be that the intercept is non-trivial (e.g. Neter et al., 1985,
pp. 163–164). At least there seems to be no valid biological reason to set b0
to zero. Inclusion of Pm as suggested above, and allowance for a non-zero
intercept in the regression, would make method 4 collapse to method 2. All
this makes method 4 unsuitable.
Method 5 has several advantages over the other methods: only a single
genetic parameter is involved and, just as in method 2, minLD/PD depends
13MechModPig 31/1/06 10:20 Page 276

276 P. Luiting and P.W.Knap

on MEI and is not constant, and b0  MEm. There are two more reasons
why this method is attractive.
First, b0 depends on P and L in such a way that b0 approaches MEm
when [minL/P × P – L] approaches zero, i.e. when L/P approaches minL/P.
This is illustrated in Fig. 13.7. When a pig that used to be fed at a high
MEI level (so that it is on a high PD metabolism, and has a high L/P level)
is suddenly brought to a low MEI level (for example, MEI = MEm as in
Fig. 13.7), it will not directly reduce its PD metabolism to zero, but will
catabolize body lipid to support its protein retention. In Fig. 13.6 this
initial situation is represented by the base of the arrow at the upper
regression line with a high L/P level (b0 < MEm). When this low MEI level
continues, the resulting lipid catabolism reduces L/P until it has reached
minL/P (through subsequently lower regression lines in Fig. 13.6), and PD
metabolism follows the arrow to approach zero as represented by the
regression line with b0 = MEm. Black et al. (1986) describe a similar effect
of b0 approaching MEm during a long period (100 days) of insufficient
energy supply. But they had to add yet another empirical equation to
method 2 to quantify this effect; method 5 does not require that.
It also follows that methods that set b0 = MEm (methods 1 and 3 in
Table 13.4) implicitly assume a steady state situation where metabolism has
been adapted to a low MEI level, another reason why they are less suitable.
Second, the concept of minL/P in method 5 is more elegant (because more
mechanistic) than the empirical approach of the linear regression
parameters in method 2. It operates on the same level as Emmans’s (1988,
1997) system of potential body protein mass and desired body lipid mass,

PD LD

maxPD

L/P > minL/P

L/P = minL/P

B

minLDmaxPD
PD A
LD

0 MEm A B C MEI




minMEI maxPD

Fig. 13.7. An extension of Fig. 13.3. The PD system is the same as in the right-hand plot of
Fig. 13. 6. Each PD line has an associated LD line which changes to a steeper slope in the
point minMEImaxPD where PD and LD reach maxPD and minLDmaxPD, respectively. See the
text for scenarios A, B and C.
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Comparison of Pig Growth Models 277

which presumes that both entities follow Gompertz functions in relation to

age, characterizsed by three genetic parameters: mature body protein mass
(Pmature), mature body lipid mass (Lmature), and a common specific growth
rate parameter BGomp.
In Emmans’s system as extended by Ferguson et al. (1994), the pig’s
desired lipid deposition at any point in time makes up for the deficit
between desired lipid mass (which follows the above genetically
determined Gompertz pattern) and actual lipid mass (which may deviate
from it for a variety of reasons). Hence for a given level of desired lipid
mass, desired lipid deposition will be lower for higher values of actual lipid
mass. Similarly, in method 5, minLDmaxPD equals [minL/P × (P + maxPD) –
L] which for a given level of minL/P is lower for higher values of L, and
hence for higher values of L/P.
In the example in Fig. 13.7, the pig with the highest actual L/P level
(farthest removed from the minL/P line) reaches its maxPD at MEI level A,
where its LD = minLDmaxPD level is lower than for the pig with the lower
actual L/P which reaches its maxPD and minLDmaxPD values at MEI level B.
The connection between Emmans’s system and method 5 can be
extended as follows.
ϕ −1
desired L L mature  P 
minL / P ≤ = ×   (13.20)
desired P Pmature  Pmature 

0.23
L 
with  = 1.46 ×  mature  as an ‘auxiliary variable’ (Emmans, 1997).
 Pmature 

More generally, the right-hand term in equation 13.20 is a non-linear

function of (P / Pmature) with genetic parameter (Lmature / Pmature). Like
maxPD it varies throughout the growth period.
It follows from the above that the two terms in inequality (equation
13.20) behave the same way, and that one serves a boundary value for the
other. This would suggest a functional relationship between the two
entities minL/P and ˇ÷ EMBED Equation.3 ˇ≠ ˇˇ§, which merits further
study.
As a consequence, minLDmaxPD must be seen as the lower limit to
desired LD. In Fig. 13.7 this leads to the situation where voluntary MEI >
minMEImaxPD so that desired LD > minLDmaxPD, indicated by the circles
for PD (at maxPD) and LD (at desired LD) when the genotype with
minMEImaxPD = B realizes a voluntary MEI = C. In that case we can write:
voluntary MEI = MEm + maxPD × EP/kP + desired LD × EL/kL =
= MEm + maxPD × EP/kP + minLDmaxPD × EL/kL + extra LD × EL/kL
(13.21)
The last term (extra LD) must then be due to either (i) a drive towards an
LD (desired LD) higher than minLDmaxPD, or (ii) a drive towards a higher
13MechModPig 31/1/06 10:20 Page 278

278 P. Luiting and P.W.Knap

MEI than minMEImaxPD. Both would lead the pig to realize a datapoint (as
in Fig. 13.7) beyond the maxPD level; Campbell et al. (1983, 1985),
Campbell and Taverner (1988), and Dunkin and Black (1987) have shown
that this actually happens. Option (ii) can more appropriately be expressed
as:
voluntary MEI − (MEm + maxPD × EP / kP + minLDmaxPD × EL / kL ) 3.20)
extra LD =
EL / kL
It follows that whereas the trajectory for MEI < minMEImaxPD is controlled by
a genetic drive for lipid deposition (expressed as minL/P), the trajectory for
MEI  minMEImaxPD is controlled by a genetic drive for protein deposition
(expressed as maxPD) and a genetic drive for either lipid deposition or
‘luxury’ ME intake (which would lead to extra LD through equation 13.22).

Final remarks

MEm is quantitatively quite large, and it depends on many environmental

factors. Hence, errors in its estimation may have considerable effects on
model predictions. All models described here assume that MEm is not
subject to genetic variation. This assumption is probably false (see Knap,
2000, p. 164) and MEm must be characterized for the genotype to be
simulated. The system of protein deposition, lipid deposition and
maintenance can be kept internally consistent if MEm and Pm are then
expressed in terms of P / Pmature (e.g. for growing animals, MEm = α ×
P
P = α × 0.73
× Pmature according to Emmans, 1997), similar to maxPD
Pmature
and desired LD (equations 13.9 and 13.18). Apart from the models
mentioned in the section on ‘Prediction of voluntary food intake’ in ‘Model
comparison’ above, most models do not simulate voluntary food intake but
treat it as an input parameter. For pig breeding applications, where
voluntary food intake is of true interest, this is unproductive; this again
requires inclusion of MEm as a genotype characteristic in the model, on the
same level as maxPD and desired LD.
Finally, all models assume EP, EL, kP and kL to be constants without
genetic variation; evidence to support or falsify this assumption is virtually
absent (see Luiting, 1990, for an overview in poultry, and Emmans, 1997).

Conclusions

For the purposes of animal breeding, desirable features of a growth model

are (i) that it is consistent with real-life data, (ii) that it is internally
consistent, (iii) that it has a small number of genotype-specific parameters,
(iv) that it is able to predict voluntary food intake. Given the shortage of
reliable data that span a sufficiently wide range of P / Pmature on modern
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Comparison of Pig Growth Models 279

genotypes, item (i) will always be hard to meet, and not necessarily as a
shortcoming of the models. For the description of maxPD, method iv of
Table 13.3 seems the most appropriate method to meet the other criteria.
With respect to feature (ii) it is best combined with method 5 of Table 13.4
for the description of PD at MEI < minMEImaxPD. An additional advantage
of this combination of methods is that it provides a smooth and consistent
connection with Taylor’s genetic size scaling principles.

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Appendix 13.1

In addition to a linear regression of PD on MEI with linear regression

coefficient r and x-intercept s, De Greef (1992) assumes a ‘constant’ linear
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Comparison of Pig Growth Models 281

regression of LD on MEI with linear regression coefficient t and x-

intercept v:
PD = r × (MEI − s)
(13.A1)
LD = t × (MEI − v)
He defines the ratio between the two linear regression coefficients as the
marginal minLD/PD ratio and assumes that this parameter is a genetically
determined constant, which makes minLD/PD dependent on MEI:
t × (MEI − v) t MEI − v MEI − v (13.A2)
minLD / PD = = × = margminLD / PD ×
r × (MEI − s) r MEI − s MEI − s
However, because of the law of conservation of energy, the linear
regression of LD on MEI can be written in terms of the parameters of the
linear regression of PD on MEI; De Greef does not mention this:
MEI − MEm − (PD × E / kP ) (13.A3)
LD = P =
EL / kL
This shows that t is a function of r, and that v is a function of r, s and MEm,
which automatically makes both t and v constants too. As a consequence,
the ratio between t and r (i.e. marg minLD/PD) is a constant with value (1 –
r × EP / kP) / (r × EL / kL).
Hence given that r = b1 and s = b0, substitution in equation 13.A2
shows that De Greef ’s minLD/PD equals the one of Black (see Table 13.2):

MEI − v
minLD / PD = margminLD / PD × =
MEI − s
MEm − r × s × EP / kP
MEI −
1 − r × EP / kP 1 − r × EP / kP
= × =
r × EL / kL MEI - s
MEm − b0 × b1 × EP / kP
MEI −
1 − b1 × EP / kP 1 − b1 × EP / kP
= × =
b1 × EL / kL MEI − b 0

 MEm − b0 × b1 × EP / kP 
MEI − b0 + b0 −  
1 − b1 × EP / kP  1 − b1 × EP / kP 
= × =
b1 × EL / kL MEI − b0
 b0 − b0 × b1 × EP / kP − MEm + b0 × b1 × EP / kP 

1 − b1 × EP / kP  1 − b1 × EP / kP 
= × 1+ =
b1 × EL / kL  MEI − b0 
 
 
1 − b1 × EP / kP  MEm − b0 1 
= × 1 − × 
b1 × EL / kL  1 − b1 × EP / kP MEI − b0  (13.A4)