Livestock Production Science 62 (1999) 113

www.elsevier.com / locate / livprodsci

On the use of simple ratios between lactation curve coefficients

to describe parity effects on milk production
a,
b
c
N.C. Friggens *, G.C. Emmans , R.F. Veerkamp
a

Department of Animal Health and Welfare, Danish Institute of Agricultural Sciences, Research Centre Foulum, P.O. Box 50,
DK-8830 Tjele, Denmark
b
Animal Biology Division, Scottish Agricultural College, West Mains Road, Edinburgh EH9 3 JG, United Kingdom
c
Department of Animal Breeding and Genetics, ID-DLO, P.O. Box 65, 8200 AB Lelystad, The Netherlands
Received 26 October 1998; received in revised form 9 April 1999; accepted 20 April 1999

Abstract
The objective of this study was to quantify how the pattern of milk production relative to time from calving is affected by
parity in cows fed high quality rations. For this purpose two models; those of Emmans and Fisher (1986) and Dijkstra et al.
(1997), were considered. Comparison with Woods (1967) function was made to evaluate their fitting ability. Daily records
of milk yield from 40 cows fed a grass silage based, high concentrate total mixed ration were used. The cows had ad libitum
access to food, they were milked twice daily. Each of the cows had milk yield records from calving to 240 days post calving
in parities 1, 2 and 3. The model of Dijkstra et al. (1997) was found, on inspection, to be an alternative parameterization of
the Emmans and Fisher model (1986). In the analyses, the Emmans and Fisher form was used: Yield 5 aU exp [2c(days
from calving)] where U 5 exp h 2 exp (G0 2 b[days from calving])j. The Emmans and Fisher model (1986) performed
marginally better than the Woods function (1967) in terms of percentage of variance accounted for and residual standard
error. There was no significant effect of parity on G0 or b, which are the main parameters describing the evolution of
lactation to peak. However, there was a highly significant effect (P , 0.001) of parity on coefficients a, the scalar, and c, the
decay coefficient. The coefficient values in parity 1 and 2 were found to be a constant proportion of the values in parity 3.
Parity effects can therefore be described by simple ratios, offering the possibility of simplifying the inputs needed in models
to describe potential milk production. 1999 Elsevier Science B.V. All rights reserved.
Keywords: Dairy cows; Lactation; Parity; Model; Milk yield

1. Introduction
The ability to predict the potential performance of
*Corresponding author. Tel.: 145-8999-1555; fax: 145-89991500.
E-mail address: n.friggens@agrsci.dk (N.C. Friggens)

dairy cows, that is the performance the cow would

achieve if it were not limited by diet or environment,
and thus calculate the energy required to support
potential, is important for two reasons. If potential
can be predicted, both relative to days in milk and
across parities, then rations can be designed that
capitalise on the genetic ability of the cow. Further,

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N.C. Friggens et al. / Livestock Production Science 62 (1999) 1 13

in models that aim to predict the partition of

nutrients between body reserves and milk production, the potential performance provides the biological basis from which to develop rules for nutrient
partitioning in all feeding situations. The largest
component of potential, in terms of energy requirements, is milk production.
It is clear that milk production varies with time,

both within and between lactations (Remond

et al.,
1997), though the major focus of studies to characterise the temporal patterns of milk production has
been on the variation within lactations. There are a
substantial number of existing lactation curve models
of varying complexity (Nelder, 1966; Wood, 1967;
Cobby and Le Du, 1978; Goodall and Sprevak, 1985;
Grossman and Koops, 1988; Morant and Gnanasakthy, 1989) and a number of studies which have
examined the performance of different models in
terms of their ability to fit milk yield data (Rowlands

et al., 1982; Rook et al., 1993; Perochon

et al., 1996)
or give accurate predictions of future yield from part
records (Wilmink, 1987; Jones, 1997). However, for
the purpose of predicting potential a high degree of
flexibility of curve shape and ability to fit environmentally or nutritionally induced distortions of the
underlying curve are not valuable attributes. The key
attributes of a useful model to describe potential are
that it is biologically interpretable and provides the
simplest sufficient description of the potential yield
curve. Both of these attributes are important when it
comes to subsequently developing modifiers to the
potential curve. Development of models to incorporate, in a logical and general manner, effects such as
those of parity, pregnancy, limiting nutrition and
environment depends upon a clear association between model parameters and the underlying biological phenomena (Dijkstra et al. 1997). The process of extending such a model to accommodate
parity effects provided the impetus for this study.
The set of lactation curve models which explicitly
aim to reflect the underlying biology is a small one
(Neal and Thornley, 1983; Emmans and Fisher,
1986; Dijkstra et al., 1997) though it can be argued
that any model which comprises explicit growth and
decay phases is biologically interpretable (Wood,
1977). All of these models, either implicitly (Dijkstra et al., 1997) or explicitly (Neal and Thornley,
1983; Emmans and Fisher, 1986) relate to potential
production.

The model of Neal and Thornley (1983) uses five

input values to define initial state, and has 14
parameters. The model of Dijkstra et al. (1997) is a
description of the rates of profileration and death of
mammary cells in simpler, and more general, terms
than the model of Neal and Thornley (1983) and can
be applied to describe the lactation curve using four
parameters. The model of Emmans and Fisher
(1986) also uses four parameters. In this model the
growth phase is described by a Gompertz function
(Gompertz, 1825; Winsor, 1932) and the declining
phase by a partial gamma decay first used for
lactation curves by Brody et al. (1923) and included
in the well established model of Wood (1967). The
same form was used by France et al. (1985) to
describe faecal marker excretion patterns.
Both the Emmans and Fisher model and the model
of Dijkstra et al. (1997) have two clear advantages
over the simpler, three-parameter, Woods function
(1967). The disadvantages of the Woods function
arise from the use of a power function, t b , to describe
the growth phase of lactation. When t 5 0, the
function predicts zero yield. The data used by Wood
(1967) were weekly records which may have made
the zero intercept seem likely. However, it is clear
from daily records that cows can have appreciable
milk yields at calving as would be expected from the
finding that the secretory potential of the mammary
gland is well developed at parturition (Knight and
Wilde, 1993). Further, t b does not have an
assymptote and so continues to have an effect,
implying continued growth of the gland, throughout
lactation. Under normal conditions there is no evidence for this (Knight and Wilde, 1993), and from
the point of view of interpreting the model, the decay
phase is not simply described but is the consequence
of two parameters. By replacing the t b term with an
asymptotic function which can have a positive
intercept, both the Emmans and Fisher model (1986)
and the model of Dijkstra et al. (1997) overcome
these two problems albeit at the cost of an extra
parameter.
Although they are derived from very different sets
of theoretical assumptions, the models of Emmans
and Fisher (1986) and Dijksta et al. (1997) would
both appear to satisfy the fundamental criteria for use
as biologically interpretable models to describe
potential milk production. However, neither of these
models, which appear to use different functional

N.C. Friggens et al. / Livestock Production Science 62 (1999) 1 13

forms, has been characterised in terms of statistical

performance. Because these models are descriptions
of potential, the appropriate data against which to
evaluate them are; milk yields from cows fed in a
manner likely to result in lactation curves whose
shapes are characteristic of potential milk production. The data used in the present study provided the
opportunity for such an evaluation.
The first objective of this study was to evaluate the
two models (Emmans and Fisher, 1986; Dijkstra et
al., 1997) in order to determine which should be
used in addressing the main objective of this study.
The ability of these models to fit milk yield data was
assessed relative to Woods function (1967).
The second, and main, objective of this study was
to quantify the effect of parity on the coefficients of
a simple, biologically based, model of the lactation
curve and, if possible, to develop simple relationships between parities in the lactation curve coefficients thus simplifying the inputs necessary for a
general model of lactational performance.

2. Materials and methods

2.1. Animals, feeding and management

Data were obtained from 40 HolsteinFriesian
cows, each of which had daily milk yield records for
first, second and third lactations. The cows were
housed and managed at the Scottish Agricultural
College / University of Edinburgh Langhill Dairy
Research Centre, Edinburgh, Scotland (longitude; 38
109 W, latitude; 558 529 N) as part of a genotype by
environment study (Veerkamp et al., 1994). Calving
occurred between September and January in any
given year. The cows were kept indoors in conventional cubicle housing from calving through to the
following July. The distribution of lactations according to calving year and parity is shown in Table 1.
The cows were milked twice daily and received 0.5
kg of a standard dairy concentrate in the milking
parlour at each milking.
All cows included in the data set were offered ad
libitum a high concentrate total mixed ration (TMR)
composed of grass silage, brewers grains and concentrate. Cows offered the low concentrate diet in
the genotype by environment study (Veerkamp et al.,

Table 1
The distribution of lactations according to calving year and parity
Parity

Calving year a
1990

1
2
3
Total

1991

1992

1993

1994

1995

8
0
0

7
8
0

13
7
8

12
13
7

0
12
13

0
0
12

15

28

32

25

12

The calving year started on the 1st September in any given

year.

1994) were not considered in the present study. The

cows were managed in three groups for feeding
according to days in lactation; less than 100 days
(early), between 100 and 200 days (mid), and greater
than 200 days (late). The proportions in total dry
matter (DM) of silage, brewers grains and concentrate in the TMRs offered to early, mid and late
lactation cows were; 40:5:55, 50:5:45 and 60:5:35
respectively. The feeding system was designed to
achieve, over a full lactation, average proportions of
silage, brewers grains and concentrate of 50:5:45 in
total DM. The average DM content (oven drying at
808C for 24 h) of the early and mid TMRs was 364
and 334 g / kg fresh, respectively. The average crude
protein content (Association of Official Analytical
Chemists, 1990) of the early and mid TMRs was 178
and 176 g / kg DM, respectively. The average
metabolisable energy content, estimated from standard ME values for the ration ingredients, of the
early and mid TMRs was 12.2 and 11.9 MJ ME / kg
DM, respectively. The average NDF content (Robertson and Van Soest, 1977) of the early and mid
TMRs was 372 and 374 g / kg DM, respectively.
Further details of the composition of the TMRs used
in the genotype by environment experiment, and of
other performance measures made, have been reported by Veerkamp et al. (1994).

2.2. Data processing

The full data set for the evaluation of the effect of
parity on lactation curve coefficients comprised
36 221 daily milk yield records. For the purpose of
deriving lactation curve coefficients, the data from
each lactation of each animal were treated as independent giving 120 lactations in all. Within each
lactation, only milk yield data from calving until 240

N.C. Friggens et al. / Livestock Production Science 62 (1999) 1 13

days post-calving were used as it has been shown

that the concurrent pregnancy in lactating dairy cows
causes a depression in milk yield in the last 18 weeks
of pregnancy (Hooper, 1923; Coulon et al., 1995).
Estimation of the lactation curve coefficients, and the
effect of parity on them, was thus unaffected by
pregnancy and drying off. Extremely deviant points
were excluded from the data in a two step process
using a spline fitting procedure with 6 degrees of
freedom (GenstatE Version 5.3.2., 1994). First, the
spline was fitted to the raw data for each lactation
curve and individual milk yields which deviated
from the spline by 65 standard deviations were
excluded from the data. Second, the spline was fitted
to the data set which excluded deviant values from
the first spline fitting. Individual milk yields which
deviated from the second spline by 65 standard
deviations were also excluded from the data. This
two step process resulted in the exclusion of, on
average, 0.98 daily milk yields per lactation, the
maximum number of data points excluded in any one
cow lactation was 6 (out of 232).

2.3. Lactation curve coefficients and statistical

analyses
To determine whether the Dijkstra et al. (1997) or
Emmans and Fisher (1986) model should be used in
addressing the main objective of this study, the first
objective of this study was to compare them in terms
of their ability to fit data. The two equations are
shown below:
dY / dt 5 a hexp f 2 exp (G0 2 bt) g j ? f exp (2ct) g
(Emmans and Fisher, 1986)
dY / dt 5 M0 ? exp h mT f 1 2 exp (2kt) g /k 2 lt j
(Dijkstra et al., 1997)
where dY / dt is milk yield per day, t is days from
calving, a, G0 , b and c are the coefficients of the
Emmans and Fisher (1986) model, and M0 , mT , k
and l are the coefficients of the Dijkstra et al. (1997)
model as stated in their Eq. (11). (The coefficient
called k in the above equation is that which Dijkstra
et al. (1997) called k 2 , the subscript has been
dropped to avoid confusion with subsequent use of
numerical subscripts in the present paper). However,

on inspection of the equation of Dijkstra et al. it

became apparent that it was an alternative form of
the model proposed by Emmans and Fisher (1986),
as shown below:
let mT /k 5 q
substituting q in the Dijkstra equation gives:
dY / dt 5 M0 ? exp f q 2 q ? exp (2kt) 2 lt g
dY / dt 5 M0 ? exp (q) ? exp f 2 q ? exp (2kt) g
? exp(2lt)
dY / dt 5 M0 ? exp (q)
? exp h 2 exp f ln (q) g ? exp (2kt) j ? exp (2lt)
dY / dt 5 M0 ? exp (q) ? exp h 2 exp f ln (q) 2 kt g j
? exp (2lt)
This is the Emmans and Fisher (1986) form
where:
M0 ? exp (q) 5 a, ln (q) 5 G0 , k 5 b and l 5 c
Given that the equation of Dijkstra et al. (1997) is
another form of the Emmans and Fisher model
(1986) it could not provide an alternative model
against which to compare the curve fit of the
Emmans and Fisher model. For this purpose, the
function of Wood (1967) was used:
dY / dt 5 at (b) ? exp(2ct)
(Wood, 1967)
where: dY / dt is milk yield per day, t is days from
calving, and a, b and c are the coefficients of the
equation of Wood (1967). The non-linear curve
fitting procedure of Genstat (GenstatE Version
5.3.2., 1994) was used to derive coefficients for the
curve of daily milk yield (dY / dt) relative to days
from calving (t) for both the Woods equation (1967)
and the equation of Emmans and Fisher (1986).
Analysis of parity effects was carried out only on
the coefficients from the Emmans and Fisher model.
The effect of parity was quantified by analysis of
variance with cow and parity within cow included as
a fixed effects (GenstatE Version 5.3.2., 1994). The

N.C. Friggens et al. / Livestock Production Science 62 (1999) 1 13

standard errors (S.E.s) of each coefficient in the

curve fitting procedure were used to weight the
analysis of variance for that coefficient by including
1 /(S.E.)2 as a covariate. Given the uneven distribution of parities across calving years (Table 1) it was
not possible to directly include calving year effects
in the analysis of variance for parity effects. Quantification of calving year effects is discussed in
Section 2.4.
Further analyses were carried out on those coefficients which were affected by parity. When
discussing these ratios, parity numbers are given as
subscripts to the relevant coefficient. For a given
coefficient, the ratio between the coefficient values
for any pair of parities (e.g., c 1 /c 3 ) was calculated
within individual cows. The average value of the
coefficient across the same pair of parities was also
calculated within individual [e.g., (c 1 1c 3 ) / 2]. The
relationship between the 40 values of the ratio
between parities and the 40 values of the average of
the coefficient over the same two parities was
examined using linear regression analyses (Minitab
Version 9.1, 1992).

2.4. Calving year effects

The effect of calving year on milk production was
examined by selecting a second data set from the
genotype by environment experiment (Veerkamp et
al., 1994). The criteria for including cows in this
second set were that; (i) they were being fed the high
concentrate TMR, (ii) they were in their third
lactation, and (iii) their third lactation was in one of
the calving years included in the parity analysis. This
resulted in a data set of 18 795 records on 67 cows,
spread evenly across calving years, which included
the third lactation cows that were in the parity
analysis. The cows in this set were managed in
exactly the same way as the cows in the parity data
set and the data were processed in the same way to
derive lactation curve coefficients for the Emmans
and Fisher model (1986). Year effects were quantified by analysis of variance with year included as a
fixed effect (GenstatE Version 5.3.2., 1994). In order
to examine if calving year would influence the parity
analysis, the average value of a given coefficient for
third lactation cows in each calving year (derived
from the calving year data set) was used as an

additional covariate in the analysis of variance for

parity effects of that coefficient (parity data set).

3. Results and discussion

The results and discussion have been presented in
an order which reflects the stepwise progress of the
analyses rather than dealing with the objectives
according to their priority. The order is: statistical
performance of the models examined and selection
of model for the parity analyses, effect of parity on
lactation curve coefficients, and finally calving year
effects.

3.1. Curve fitting

The average lactation curves for each parity are
shown in Fig. 1. One cow was removed from the
study because of chronic health problems in two (out
of three) of her lactations. An additional six lactations, out of 117, were discarded because of chronic
health problems. Using the Emmans and Fisher
model (1986) the curve fitting procedure failed to
converge in a further six cases, this was due to runs
of missing data at the start of lactation for these
particular lactations. For the 105 cases where the
curve fitting procedure converged, the average R 2 of
the curve fit was 84.1% (min.545.0; first quartile5
80.4; third quartile591.3; max.597.8%) and the
average residual standard deviation of the curve fit
was 1.74 kg / d (min.50.76; first quartile51.38; third
quartile52.02; max.53.89 kg / d). The residuals
from the curve fit of the average milk yield curve of
third lactation cows (Fig. 1) are shown for the
Emmans and Fisher model (1986) in Fig. 2. The
correlations between the coefficients a, G0 , b and c,
of the Emmans and Fisher model (1986) are shown
in Table 2. The relationship between coefficients a
and c of the Emmans and Fisher model (1986) is
shown in Fig. 3.
For the 105 cases where curve fit statistics were
available for both models, the average R 2 of the
Woods curve (1967) fit was 82.1% (min.530.1, first
quartile579.1, third quartile590.4, max.597.8%)
and the average residual standard deviation of the
Woods curve fit was 1.86 kg / d (min.50.82, first
quartile51.42, third quartile52.15, max.53.78 kg /

N.C. Friggens et al. / Livestock Production Science 62 (1999) 1 13

Fig. 1. The relationship between average milk yield and time from calving for cows in parity 1, 2 and 3. The curve with the highest peak
yield is parity 3, the curve with the lowest peak yield is parity 1.

Fig. 2. The residuals from non-linear regression of daily average milk yield of third lactation cows relative to days post calving using: (d)
the Emmans and Fisher model (1986), and (s) Woods function (1967). The fitted curve (Emmans and Fisher, 1986) is shown for reference
(solid line).

d). The curve fitting procedure converged in all cases

using the equation of Wood (1967) indicating that it
was more robust, in these terms, than the Emmans
and Fisher model (1986). This is not surprising given

that the Woods function has a fixed zero intercept at

calving and was thus not vulnerable to runs of
missing data in early lactation. Around peak yield
and in mid lactation, there was a tendency for greater

N.C. Friggens et al. / Livestock Production Science 62 (1999) 1 13

Table 2
The correlations between the coefficients of the Emmans and
Fisher (1986) lactation curve model, calculated in two ways a
G0
G0
b
a
c

b
0.625

0.424
0.098
0.196

20.269
20.199

20.117
20.686

20.063
20.612
0.937

0.750

Within each of the 105 cow lactations where the curve fitting
procedure converged, the correlation between coefficients was
estimated in the curve fit. Each value above the diagonal is the
average of the corresponding 105 correlations. Below the diagonal, each value is a single correlation, across cows and lactations,
between the 105 values of the relevant two coefficients. There was
no significant effect of parity on the correlations.

deviations from zero in the residuals from the curve

fitting procedure when using the Woods function
relative to the Emmans and Fisher model (Fig. 2).
On the basis of the curve fit statistics presented
above there is relatively little to distinguish between
the models of Wood (1967) and Emmans and Fisher
(1986) but, as discussed in Section 1, the Emmans
and Fisher model gives a better representation of the
onset of lactation and is more easily interpretable in
biological terms.
The above is, to our knowledge, the first published
account of the ability of the Emmans and Fisher
model to fit the milk yield of dairy cows whose
lactation curves can reasonably be assumed to be

representative of potential. Rook et al. (1993) compared a range of lactation curve models including
one which was a form of the Emmans and Fisher
model. However, this study (Rook et al., 1993) is not
directly relevant to the present study as the data used
were derived from cows which for the most part
were fed in a way likely to result in nutrition
distortions of lactation curves relative to potential
milk production.
In the study of Rook et al. (1993) their form of the
Emmans and Fisher model [Eq. (7A); Rook et al.,
1993] did not perform well, ranking 11th out of 13
models (on the basis of mean rank of residual mean
squares). The Woods function (1967) ranked third
out of the 13 models in their study (Rook et al.,
1993). In contrast, in the present study the Emmans
and Fisher model (1986) performed marginally
better than the Woods function (1967) in terms of
percentage of variance accounted for and residual
standard error. In our view, the difference found
between the present study and that of Rook et al.
(1993) in the statistical performance of the Emmans
and Fisher model (relative to Woods function)
serves to emphasis the importance of using suitable
data for testing models of potential production.
The original reason for comparing lactation curve
models in this study was to select for the subsequent
parity analyses the better performing of two biologically interpretable models (Emmans and Fisher,

Fig. 3. The relationship between coefficients a and c of the Emmans and Fisher lactation curve model.

N.C. Friggens et al. / Livestock Production Science 62 (1999) 1 13

1986; Dijkstra et al., 1997). These models were

derived from substantially different perspectives and
there appeared to be no obvious equivalence between
them. However, as shown in Section 2.3, the model
of Dijkstra et al. (1997) in fact reduces to the earlier
Emmans and Fisher model (1986). This applies to
the equation of Dijkstra et al. (1997) for predicting
milk yield [Eq. (11); Dijkstra et al., 1997] and to the
equation for mammary growth post-partum [Eq.
(5b); Dijkstra et al., 1997] because their parameter T
is a constant within species. The comments do not
relate to the pre-partum mammary growth model
(Dijkstra et al., 1997). The fact that the same model
arises from two distinctly different philosophies for
describing the biological processes or mechanisms
underlying milk production is reassuring. Indeed, one
basis for ascribing generality to a functional form or
theory is that it has arisen independently from
different approaches or disciplines (Von Bertalanffy,
1968).
The choice of which parameterisation to use
[Emmans and Fisher (1986) or Dijkstra et al. (1997)]
is dependent upon how well the parameter interpretations relate to the purpose for which the model is to
be used. The mathematical equivalence between the
two parmeterisations is given in Section 2.3. In
biological terms, the roles of the different parameters
can be summarised thus.
The scaling coefficient, a, in the Emmans and
Fisher model (1986) can be seen as the main
coefficient by which differences between cows in
milk yield potential would be expressed (Congleton,
Jr. and Everett, 1980). It does not, however, directly
give milk yield at any time in lactation and has no
direct equivalence to any one parameter in the
equation of Dijkstra et al. (1997). The coefficient
which allows scaling in the equation of Dijkstra et al.
(1997) is M0 .
The growth of the milk producing system is
determined by G0 and b in the Emmans and Fisher
model (1986), and by mT and k in the equation of
Dijkstra et al. (1997). The rate at which milk yield
increases to peak is described by a single coefficient
in both models, b (Emmans and Fisher, 1986) and k
(Dijkstra et al. 1997). The coefficient G0 in the
Emmans and Fisher model (1986) quantifies the
degree of maturity of the milk producing system at
calving through the expression exp [2exp (G0 )]

which is bounded between 0 and 1 (when G0 tends to

2` and `, respectively). The average value of G0 in
the present study indicates that the milk producing
system is approximately 40% developed at calving.
The coefficient M0 in the equation of Dijkstra et al.
(1997) is equal to the product of the Emmans and
Fisher milk yield scalar, a, and their degree of
maturity expression, exp [2exp (G0 )]. Consequently,
M0 gives the milk yield at calving which in this
study was calculated as (kg / d); 14.2, 20.5 and 21.2
for parities 1, 2 and 3, respectively. The coefficient
mT in the equation of Dijkstra et al. (1997) is defined
as the rate of secretory cell proliferation at calving
(Dijkstra et al., 1997) and thus relates to descriptions
of mammary growth but does not easily translate to a
describable property of the milk production curve.
For a given value of k, mT controls the amplitude of
the growth phase of the milk yield curve but said
amplitude is expressed in a ratio with k and as an
exponential multiplier for M0 [M0 ?exp ( mT /k)].
In both the model of Emmans and Fisher (1986)
and the equation of Dijkstra et al. (1997) the rate of
decline in milk yield is described by a single
coefficient, called c and l, respectively. In terms of
biological interpretation this has a clear advantage
over the Woods function (1967) where the rate of
decline is controlled by two parameters. There is,
however, a good argument for introducing a second
parameter which relates to the declining phase of
lactation, to describe the effect of pregnancy in
depressing concurrent milk production (Hooper,
1923; Coulon et al., 1995). Such modifiers have been
proposed (Coulon et al., 1995) and indeed parameter
b in the Woods function (1967) may provide the
flexibility to account for the depression in milk yield
due to the extent of pregnancy in late lactation.
However, in order to be biologically sensible, a
parameter to describe the effect of pregnancy should
clearly be related to the date of conception of the calf
and be independent of the stage of lactation of the
cow.

3.2. Parity effects

The results concerning both parity effects and
calving year relate only to the Emmans and Fisher
model (1986). The average values of the lactation
curve coefficients in parities 1, 2 and 3 are presented

N.C. Friggens et al. / Livestock Production Science 62 (1999) 1 13

Table 3
The average values of the coefficients of the Emmans and Fisher
(1986) lactation curve model for parities 1, 2 and 3
Parity

Coefficient
G0

1
2
3

20.206
20.245
20.089

0.0694
0.0916
0.0888

32.1
44.9
52.8

0.00218
0.00322
0.00393

SED a
Pb

0.072
ns

0.0118
ns

1.7
***

0.00023
***

Standard error of the difference.

Significance of the effect of parity: ns indicates P.0.05, ***
indicates P,0.001.
b

in Table 3. There were no significant effects of parity

on coefficients b and G0 (Table 3). This indicates
that any effects of parity on both the degree of
maturity of the milk producing system (G0 ) and the
rate at which milk production increases to peak (b)
are small relative to the variation present. Thus,
when generating potential milk yield curves for
different parities in general prediction models, a
single average value across parities of both b and G0
can be used. However, there were highly significant
effects of parity on coefficients a and c (P,0.001;
Table 3). The lactation curve scalar, a, increased
indicating that potential yields increased with increasing parity. The decay coefficient, c, also in-

creased indicating that the rate of decline in milk

yield post-peak was progressively steeper with increasing parity. Similar significant effects of parity
on Woods function (1967) have been frequently
reported (Rao and Sundaresan, 1979; Congleton Jr.
and Everett, 1980; Wood, 1980; Rowlands et al.,
1982; Yadav and Sharma, 1985; Collins-Lusweti,
1991). However, and in contrast to the above results,
as the coefficients of Woods function are not
mutually exclusive in describing the underlying
biology they are all affected by parity.
The finding that only two of the lactation curve
coefficients were affected by parity suggests that a
simplification in the inputs needed to generate lactation curves for different parities may be possible.
This depends upon the relationship between the
coefficients of the Emmans and Fisher model (1986)
across parities. The data used in the study of parity
effects were chosen so as to allow this relationship to
be examined. For each cow, the ratio in the value of
coefficient a between parities 1 and 2, 1 and 3, and 2
and 3 was calculated. The same ratios were also
calculated for coefficient c. The averages, across
cows, of these ratios are presented in Table 4. In
order to test whether the values of these ratios were
independent of the size of the coefficient, regression
analyses between the ratios and the average values of
the coefficient were carried out. Examples of the
relationship between the ratio and the average value

Table 4
The average values of the ratios, calculated within individuals, between parities for coefficients a and c of the Emmans and Fisher model
(1986)a
Ratio between parities b
a 1 /a 2
Mean
S.E.M.c

0.73
0.029

Residual SD d
R 2 (%)d
Slope coefficient d
S.E. of slope d
Pd

0.163
4.6
0.0075
0.00474
ns

a 1 /a 3

a 2 /a 3

0.63
0.032

0.88
0.030

0.177
0.0
20.0014
0.00488
ns

0.168
3.7
20.0059
0.00401
ns

c 1 /c 2

c 1 /c 3

c 2 /c 3

0.69
0.067

0.57
0.054

0.87
0.048

0.348
17.4
168
60.3
**

0.276
10.3
86
42.0
ns

0.276
0.0
226
38.9
ns

Summary statistics are also presented for linear regressions, across cows, of the ratios (e.g., a 1 /a 2 ) on the averages of the coefficients in
the same ratio [e.g., (a 1 1a 2 ) / 2].
b
Parities are indicated by the subscript numbers in the ratios.
c
Standard error of the mean.
d
Summary statistics from the regression analyses: R 2 is the percentage of variance accounted for by the regression, adjusted for degrees
of freedom; P is the significance of the slope coefficient, ns indicates P.0.05, ** indicates P.0.01.

10

N.C. Friggens et al. / Livestock Production Science 62 (1999) 1 13

Fig. 4. The ratio of the value of coefficient a (Emmans and Fisher, 1986) in parity 1 (a 1 ) and parity 3 (a 3 ) shown relative to the average
value of the coefficient across these two parities [(a 1 1a 3 ) / 2]. Both the ratios and the averages were calculated within cows. The line shows
the average value of the ratio.

of the coefficient are shown for coefficient a in Fig.

4 and coefficient c in Fig. 5. With one exception, the
slopes of all the regressions were not significantly
different from zero and consequently the proportion
of variance accounted for by the regressions was
very low (Table 4). This indicates that for both
coefficient a and coefficient c the ratios were independent of the average size of the coefficient. The
exception was the ratio between parities 1 and 2 of
coefficient c which is shown in Fig. 5. The regression equation of the relationship between the ratio
and the average value of c across these two parities
was: c 1 /c 2 50.2471168[(c 1 1c 2 ) / 2]. However, the
regression accounted for only 17.4% of the total
variation.
There was a significant slope in only one out of
six regressions. Given that this relationship had a
correlation coefficient of 0.42, it does not provide
reasonable grounds for rejecting the simpler finding
that the ratios between parities in coefficients a and c
were independent of the average size of the coefficient. Thus, in the present study, the differences
between parity in the shape of the lactation curve
were accounted for, across cows, by simple stable
ratios in the coefficients a and c. This offers a

general means to generate potential milk yield curves

for different parities from information relating to one
parity only. Accounting for the proportion of the
variation due to the general effect of parity, independent of coefficient size, is also important
because it allows true individual variation to be
quantified and exploited (Taylor, 1985). However,
this represents an issue subsequent to and outside the
scope of the present study.
Further, if an assumption is made about the
relationship between coefficients a and c (Emmans
and Fisher, 1986) across cows within the chosen
reference parity then potential milk yield curves can
be generated for all parities from one single value
such as desired or potential 305 day yield. There is
evidence to suggest that the main difference between
high and low producing cows (of equal parity) in the
shape of the lactation curve is in the scaling parameter, a, (Congleton Jr. and Everett, 1980). Thus, for
the purpose of predicting potential lactation curves, it
may be reasonable to assume that coefficient c is a
constant within parity, independent of a. An alternative assumption could make use of the relation
between a and c shown in Fig. 3 which can be
described by linear regression as: a525.015918(c)

N.C. Friggens et al. / Livestock Production Science 62 (1999) 1 13

11

Fig. 5. The ratio of the value of coefficient c (Emmans and Fisher, 1986) in parity 1 (c 1 ) and parity 2 (c 2 ) shown relative to the average
value of the coefficient across these two parities [(c 1 1c 2 ) / 2]. Both the ratios and the averages were calculated within cows. The line shows
the average value of the ratio.

(R 2 555.8%, residual SD58.04 kg). This may be a

reasonable assumption for the purposes of prediction
but it clearly is due, in large part, to the form of the
lactation model and the associated correlations in the
coefficient estimates (Table 2).

3.3. Calving year effects

The data set used for the parity analysis was
chosen so as to be able to obtain estimates of the
ratios between parities for lactation curve coefficients
that were not biased by variation between cows; the
same cows were present in each parity. The inevitable consequence of this approach was that there was
an uneven distribution of parities across calving
years with more heifers present in the early years and
more third lactation cows in the later years (Table 1).
Thus, the design of the experiment was vulnerable to
both systematic effects of calving year on lactational
performance and extreme effects in the first and last
years. To quantify whether such calving year effects
existed a second data set was constructed, as described in Section 2.4. The average values of the
coefficients of the Emmans and Fisher model (1986)
for each calving year are presented in Table 5. There

Table 5
The average values of the coefficients of the Emmans and Fisher
model, derived from a control data set of third parity cows, for
each calving year included in the parity analysis
Calving year a

Coefficient
G0

1990
1991
1992
1993
1994
1995

0.456
0.079
0.074
20.177
0.089
20.041

0.0967
0.0993
0.0784
0.0961
0.0932
0.0872

51.4
53.2
74.1
53.1
60.3
52.2

0.00399
0.00374
0.00566
0.00341
0.00479
0.00366

Overall mean
SED b
Pc

0.0608
0.1817
ns

0.0918
0.03056
ns

57.1
14.13
ns

0.00417
0.00023
ns

The calving year started on the 1st September in any given

year.
b
Standard error of the difference.
c
Significance of the effect of year: ns indicates P.0.05.

were no significant effects of calving year on any of

the four coefficients. Further, the average values of a
given coefficient for each calving year (Table 5)
were used as a covariate in the analysis of variance
for the effect of parity on the same coefficient in the

12

N.C. Friggens et al. / Livestock Production Science 62 (1999) 1 13

parity data set. In all cases, inclusion of the calving

year effect covariate had no significant effect on the
analysis for parity effects. These results indicate that
the analysis of the effects of parity on the lactation
curve coefficients and the calculation of the ratios
between parities for the curve coefficients were not
affected by the uneven distribution of parities across
calving years.

3.4. Further considerations

The main aim of this study, to provide a simple
means by which to generate potential lactation
curves for different parities from limited information,
has been achieved. This has been done using data
from cows managed within one system, in one herd.
Thus the results presented here can be seen as a first
step, an important subsequent step would be to
examine these relationships in other herds which
have been managed and fed in a manner likely to
result in milk yield curves whose shapes are characteristic of potential milk production. We have been
careful to present our study in terms of describing
potential, the reason for this being that both the aims
and the model used in the study were not designed to
be able to accommodate the effects of sub-optimal
feeding on milk production. Models to do this clearly
require to consider the balance of nutrient inputs and
outputs as well as the potential of the animal
(Oldham and Emmans, 1989). We therefore chose to
study cows in an experiment in which it could be
reasonably assumed that the resulting lactation
curves reflected potential. However, no single experiment can be sure of having observed potential,
providing a further reason to examine these relationships in other herds. We hope that the issues raised in
this study will be taken up by others and further
developed.

4. Conclusions
The Emmans and Fisher model (1986) provides a
simple, biologically interpretable, description of the
potential yield curve that overcomes the limitations
of the functional form of the Woods function
(1967). In terms of statistical performance, the

Emmans and Fisher model (1986) was, if anything,

marginally better than the Woods function (1967).
Parity had significant effects on only two of the
coefficients of the Emmans and Fisher model (1986),
a and c. The effect of parity was adequately described by simple ratios between any pair of parities
for a given coefficient. Thus, the coefficient values in
parity 1 and 2 were found to be a constant proportion
of the values in parity 3. This allows the general
effect of parity to be accounted for, both in models
to predict potential milk yield and in analyses to
characterise variation between individual cows.

Acknowledgements
The technical assistance of the staff at the Langhill
Dairy Research Centre (Edinburgh, Scotland, UK) is
of the
gratefully acknowledged. Inge Riis Korsgard
Biometry and Genetics Department, Danish Institute
of Agricultural Sciences provided valuable statistical
help for which we are grateful. This study was
funded by the Ministry of Agriculture Fisheries and
Food (Consortium DS04; RUMINT project). The data
used were collected as part of a project funded by the
Scottish Office Agriculture, Environment and
Fisheries Department, the Milk Marketing Board for
England and Wales, the Holstein Friesian Society for
Great Britain and Ireland, and the Ministry of
Agriculture Fisheries and Food.

References
Association of Official Analytical Chemists, 1990. Official Methods of Analysis, AOAC, Washington, DC.
Brody, S., Ragsdale, A.C., Turner, C.W., 1923. The rate of decline
of milk secretion with the advance of the period of lactation. J.
Gen. Physiol. 5, 441444.
Cobby, J.M., Le Du, Y.L.P., 1978. On fitting curves to lactation
data. Anim. Prod. 26, 127133.
Collins-Lusweti, E., 1991. Lactation curves of HolsteinFriesian
and Jersey cows in Zimbabwe. S. Afr. J. Anim. Sci. 21, 1115.
Congleton, Jr. W.R., Everett, R.W., 1980. Application of the
incomplete gamma function to predict cumulative milk production. J. Dairy Sci. 63, 109119.

Coulon, J.B., Perochon,

L., Lescourret, F., 1995. Modelling the
effect of the stage of pregnancy on dairy cows milk yield.
Anim. Sci. 60, 401408.
Dijkstra, J., France, J., Dhanoa, M.S., Maas, J.A., Hanigan, M.D.,

N.C. Friggens et al. / Livestock Production Science 62 (1999) 1 13

Rook, A.J., Beever, D.E., 1997. A model to describe growth
patterns of the mammary gland during pregnancy and lactation.
J. Dairy Sci. 80, 23402354.
Emmans, G.C., Fisher, C., 1986. Problems in nutritional theory.
In: Fisher, C., Boorman, K.N. (Eds.), Nutrient Requirements of
Poultry and Nutritional Research, Butterworths, London, pp.
939.
France, J., Thornley, J.H.M., Dhanoa, M.S., Siddons, R.C., 1985.
On the mathematics of digesta flow kinetics. J. Theor. Biol.
113, 743748.
Genstat Version 5.3.2, Clarendon Press, Oxford.
Gompertz, B., 1825. On the nature of the function expressive of
the law of human mortality and on a new method of determining the value of life contingencies. Phil. Trans. Royal Soc. 115,
513585.
Goodall, E.A., Sprevak, D.A., 1985. A Bayesian estimation of the
lactation curve. Anim. Prod. 40, 189194.
Grossman, M., Koops, W.J., 1988. Multiphasic analysis of lactation curves in dairy cattle. J. Dairy Sci. 71, 15981608.
Hooper, J.J., 1923. Studies of dairy cattle. II. Milk production.
Kentucky Agric. Exp. Station Bull. 248, 6585.
Jones, T., 1997. Empirical bayes prediction of 305-day milk
production. J. Dairy Sci. 80, 10601075.
Knight, C.H., Wilde, C.J., 1993. Mammary cell changes during
pregnancy and lactation. Livest. Prod. Sci. 35, 319.
Minitab Inc, 1992. Minitab Version 9.1, Minitab Inc, State
College, PA.
Morant, S.V., Gnanasakthy, A., 1989. A new approach to the
mathematical formulation of lactation curves. Anim. Prod. 49,
151162.
Neal, H.D.S.C., Thornley, J.H.M., 1983. The lactation curve in
cattle: a mathematical model of the mammary gland. J. Agric.
Sci. Camb. 101, 389400.
Nelder, J.A., 1966. Inverse polynomials, a useful group of multifactor response functions. Biometrics 22, 128141.
Oldham, J.D., Emmans, G.C., 1989. Prediction of responses to
required nutrients in dairy cows. J. Dairy Sci. 72, 32123229.

Perochon,
L., Coulon, J.B., Lescourret, F., 1996. Modelling
lactation curves of dairy cows with emphasis on individual
variability. Anim. Sci. 63, 189200.

13

Rao, M.K., Sundaresan, D., 1979. Influence of environment and

heredity on the shape of lactation curves in Sahiwal cows. J.
Agric. Sci. 92, 393401.

Remond,
B., Rouel, J., Pinson, N., Jabet, S., 1997. An attempt to
omit the dry period over three consecutive lactations in dairy
cows. Ann. Zootech. 46, 399408.
Robertson, J.B., Van Soest, P.J., 1977. Dietary fiber estimation in
concentrate animal feed-stuffs. 69th Meeting of the American
Society of Animal Science. J. Anim. Sci. 54 (Suppl. 1),
254255.
Rook, A.J., France, J., Dhanoa, M.S., 1993. On the mathematical
description of lactation curves. J. Agric. Sci. Camb. 121,
97102.
Rowlands, G.J., Lucey, S., Russell, A.M., 1982. A comparison of
different models of the lactation curve in dairy cattle. Anim.
Prod. 35, 135144.
Taylor, St. C.S., 1985. Use of genetic size-scaling in evaluation of
animal growth. J. Anim. Sci. 61 (Suppl. 2), 118143.
Veerkamp, R.F., Simm, G., Oldham, J.D., 1994. Effects of
interaction between genotype and feeding system on milk
production, feed intake, efficiency and body tissue mobilization
in dairy cows. Livest. Prod. Sci. 39, 229241.
Von Bertalanffy, L., 1968. General System Theory, George
Braziller, New York.
Wilmink, J.B.M., 1987. Comparison of different methods of
predicting 305-day milk yield using means calculated from
within-herd lactation curves. Livest. Prod. Sci. 17, 117.
Winsor, C.P., 1932. The Gompertz curve as a growth curve. Proc.
Natl. Acad. Sci. 18, 18.
Wood, P.D.P., 1967. Algebraic model of the lactation curve in
cattle. Nature 216, 164165.
Wood, P.D.P., 1977. The biometry of lactation. J. Agric. Sci.
Camb. 88, 333339.
Wood, P.D.P., 1980. Breed variations in the shape of the lactation
curve of cattle and their implications for efficiency. Anim.
Prod. 31, 131141.
Yadav, S.B.S., Sharma, J.S., 1985. Functions for lactation curves
in crossbred dairy cattle. Indian J. Anim. Sci. 55, 4247.