A Model of Diurnal Grazing Patterns and Herbage Intake of A Dairy Cow, Mindy Model Description

Ecological Modelling 270 (2013) 1129
Contents lists available at ScienceDirect
Ecological Modelling
journal homepage: www.elsevier.com/locate/ecolmodel
A model of diurnal grazing patterns and herbage intake of a dairy cow,

MINDY: Model description
Pablo Gregorini a, , Pierre C. Beukes a , Alvaro J. Romera a , Gil Levy a , Mark D. Hanigan b
a
b
DairyNZ, Ltd., Private Bag 3221, Hamilton, New Zealand

Virginia Polytechnic Institute and State University, Blacksburg, VA 24061, USA
a r t i c l e
i n f o
Article history:
Received 23 June 2013
Received in revised form 28 August 2013
Accepted 3 September 2013
Available online 5 October 2013
Keywords:
Modeling
Grazing behavior
Herbage intake
Cattle
a b s t r a c t
Estimates of herbage intake and parallel measurements of ingestive and digestive behaviors of grazing ruminants pose considerable experimental and technical difculties, owing to dynamic interactions
between the plant, the rumen and the animal. As a consequence, advances in the area have been slow
and costly. Model simulations that capture such interactions are critical for research and management
decisions involving the grazing process. This work describes MINDY, a mathematical, mechanistic and
dynamic simulation model of the diurnal grazing pattern of a dairy cow. MINDY is based on a cluster of
three models: (1) Molly (Baldwin, 1995), a model of ruminant digestion and metabolism; (2) a model
representing feed consumption as a function of diurnal uctuations in the internal state of the animal;
and (3) a sward structure, herbage quality and grazing behavior model. The objective of the work was
to describe the diurnal grazing pattern, including ingestive actions and rumination behaviors, herbage
intake, and nutrient supply to the animal in response to the animals internal state and grazing environment. The model was coded in ACSL and simulations were conducted using ACSLXtreme. In addition
to dietary nutrient composition required by Molly, MINDY requires sward surface height and mass, and
grazing area offered to the cow. Key sub-model parameters were identied by sensitivity analyses and
parameterized using two data sets from mid-lactation Friesian and late lactation Holstein dairy cows
breeds under set stock conditions. The parameterized model predicted realistic estimates of ingestive
behavior for different cow genotypes managed under set stocking and rotational grazing. It also predicted a realistic number of steps taken while eating and searching and sward defoliation dynamics as
well as diurnal uctuations of digestion and metabolism. Additional evaluations are required and further
data may be needed to better dene some parameters, but the model offers promise as an heuristic tool
for feed intake and grazing process research and as an informative tool for grazing and cow management
decisions.
2013 Elsevier B.V. All rights reserved.
1. Introduction
Herbage intake is the most important variable affecting animal
performance in pastoral production systems (Kolver and Muller,
1998; Burns and Sollenberger, 2002). No reliable means exist to
measure it, therefore, its prediction and modeling are critical for
supporting targeted research to improve management of intake
in grazing systems. Herbage intake results from grazing (Gibb,
1996), a complex process involving animal decisions at multiple
spatio-temporal scales (Arnold and Dudzinski, 1978; Senft et al.,
1987; Bailey et al., 1996). Considerable research has been conducted on several aspects of this process, from bite formation to
Corresponding author. Tel.: +64 7 858 3787; fax: +64 7 858 375.
E-mail addresses: Pablo.Gregorini@dairynz.co.nz (P. Gregorini),
Pierre.Beukes@dairynz.co.nz (P.C. Beukes), Alvaro.Romera@dairynz.co.nz
(A.J. Romera), Gil.Levy@dairynz.co.nz (G. Levy), mhanigan@vt.edu (M.D. Hanigan).
0304-3800/$ see front matter 2013 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.ecolmodel.2013.09.001
long-term digestive constraints and grazing distribution patterns

in a landscape (Heitschmidt and Stuth, 1991; Jung and Fahey, 1999;
Gregorini, 2012), and a large body of empirical data has been accumulated.
Modeling has focused on the mechanics and dynamics of the
grazing process. Ungar and Noy-Meir (1988) and Parsons et al.
(1994) developed model to relate intake rate to herbage availability, sward structure, and leaf area index of two contrasting plant
species. Baker et al. (1992) and Baumont et al. (2004) developed
more complex models based on sub-models of grazing behavior sward structure and the animal. Brereton et al. (2005) and
Woodward (1998) modeled herbage utilization by cattle for rotational grazing systems. In all cases however, the animal component
of these models was elementary, which constraints detailed representations of the effect of animal metabolism and physiological
state on feeding decisions. Recently, Chilibroste et al. (2008) developed a simulation model to predict nutrient supply to a dairy cow
under discontinuous feeding patterns. Regardless of complexity,
12
P. Gregorini et al. / Ecological Modelling 270 (2013) 1129
Fig. 1. Schematic representation of MINDY: a mechanistic and dynamic model to simulate diurnal patterns of herbage intake and grazing behavior of a grazing dairy cow.
White boxes with solid lines represent true pools (hard components) of the model, white dashed with dashed lines represent soft components of the model, solid arrows
represent modiers. Grey boxes (functional components) and arrows represent the motivational system of feeding behavior adapted from Jensen and Toates (1993), Hughes
and Duncan (1988) and Smith (1996).
none of these models dynamically integrate animal internal state

with spatio-temporal uctuations in sward structure and herbage
chemical composition. Nor they account for photoperiod effects
on diurnal, daily and seasonal changes in feeding motivation and
intake for a grazing dairy cow.
Animal behavior is controlled by stimulation systems (Smith,
1996; Toates, 2002) that motivate certain responses (Day et al.,
1998), e.g. grazing, which is dened as the process of seeking, harvesting and orally processing herbage before swallowing (Gibb,
1996). Two types of stimulation systems motivate grazing: hunger
and incentives (Staddon, 1983; Tolkamp et al., 1998). The former
results from integration of direct and short-term nutritive (e.g.
rumen function) and physiological stimuli (e.g. hormone release)
modulated by long-term stimuli (e.g. physiological state and adiposity) (Kissileff and Van Itallie, 1982; Weston, 1996; Geary, 2004).
The incentives are an integration of the animals internal state (e.g.
hunger) in the context of the grazing environment, what establishes a functional relationship between direct and indirect control
of feeding motivation (Kyriazakis et al., 1999).
Spatio-temporal variability in sward characteristics means that
grazing environments are complex by nature. The sward canopy
(structure, morphology and chemical composition) changes in
space (horizontal and vertical planes) and time. These changes
inuence availability and accessibility of herbage and thereby grazing (Drescher, 2003; Thompson Hobbs et al., 2003). Ruminants eat
in periodic and discrete meals, which add to total daily herbage
intake (Gibb, 1996). Hence, behavioral decisions such as when to
begin and end a meal, meal frequency, meal patterns within a day,
and harvesting intensity within each meal (cumulatively dened as
the diurnal grazing pattern) determine how animals allocate time
to feeding and thereby, the form and rate of nutrient supply for
digestion, metabolism and growth (Gregorini, 2012).
The primary objective of the work reported here was to model
the diurnal patterns of ingestive and digestive behavior (including rumination), the resulting herbage intake and nutrient supply
in response to the interaction between the animal internal state
and the grazing environment of a dairy cow. This work extends the
efforts of Baldwin (1995), Baker et al. (1992), Galli et al. (1999)
and Baumont et al. (2004). The mathematical formulation, rst
principles and model parameters are described in this paper. For

illustrative purposes, some model outputs are presented using an
old and a modern New Zealand HolsteinFriesian biotype cow
grazed under set stocking and rotational grazing (24 h pasture
strips) methods on a Lolium perenne L. dominated sward. The model
has wider application. Comparisons between experimental data
and simulated values will be presented in subsequent manuscripts,
as well as a description of a supplementation and a time at pasture
restrictions scheme.
2. Model description: MINDY
MINDY is a mechanistic and dynamic simulation model of the
diurnal grazing pattern of a dairy cow (Fig. 1). MINDY is based on a
cluster of three models: (1) the dairy cow digestion and metabolism
model of Baldwin (1995) modied by Hanigan et al. (2013) and
known as Molly; (2) a model of diurnal uctuations in feeding motivation; and (3) a model of sward canopy structure, herbage quality
and grazing behavior. The latter were derived from the models of
Baumont et al. (2004) and Galli et al. (1999).
A glossary of parameters and variable names, parameter values, and denitions is included in the Appendix (Table A.1). Code
was developed and simulations were conducted using ACSLXtreme
(Ver. 2.5.0.6 Aegis Technologies Group, Austin, TX). Numerical integration was conducted using a fourth-order, xed-step,
RungeKutta method. The maximum integration interval was set
to 1 min. Results were collected after 8 d of simulation to ensure
the model had reached a stable state.
2.1. Internal state: rumen digestion, metabolism and lactation
The internal state of the animal (internal stimulation systems,
Fig. 1) and productive parameters are represented using Molly
(Baldwin, 1995) with recent modications to improve the accuracy
prediction of lipid, glucose and energy metabolism (McNamara and
Baldwin, 2000), lactation curves (Palliser and Woodward, 2002),
photoperiod effects on milk production (Beukes et al., 2005), lactation potential (Hanigan et al., 2008) and anabolic and catabolic
hormone dynamics, as well as gestational metabolism (Hanigan
13
Fig. 2. A simplied schematic representation of Baldwin (1995). Adapted from Hanigan et al. (2008). Boxes with dashes lines represent conceptual compartments as dened
in the model; boxes with solid lines represent pools; solid arrows represent modiers; and the circles associated with dashed arrows indicate the direction of the modies.
FA, fatty acids, VFA, volatile fatty acids.
et al., 2009) of dairy cows under grazing conditions. Recently the

digestive elements of the model were reparameterized using a
newer and broader data set and several representations of rumen
function were updated (Hanigan et al., 2013). An overview of that
model is depicted in Fig. 2. For details of this model, the reader is
referred to Baldwin (1995) and the subsequent developments of
Molly (Hanigan et al., 2008, 2009, 2013; Gregorini et al., 2013).
2.2. Diurnal uctuations of feeding motivation
2.2.1. Photoperiod
The diurnal timing of meals differs distinctly among animal
species (Collier and Johnson, 1990). In grazing ruminants the meal
and ingestive behavior patterns are circadian (Gregorini, 2012).
Grazing during daylight hours is highly preferred (Rook and Huckle,
1997; Linnane et al., 2001), although short grazing bouts (515% of
daily grazing time) may occur during the night (Krysl and Hess,
1993). In temperate climates, ruminants have three to ve major
meals per day (Hodgson, 1990; Gibb et al., 1998). This frequency
is exible and affected by external stimuli (e.g. photoperiod) and
behavioral adaptations to the grazing management and sward state
(Gibb, 1996; Gregorini, 2012). For example, during short days,
meals merge and the number of daily meals decreases, probably in an attempt to maximize grazing time during light hours
(Rook and Huckle, 1997). Regardless of frequency, the major meals
occur around twilight hours, dawn and dusk, with the latter having greater intensity and longer duration (Gibb et al., 1998; Taweel
et al., 2004; Gregorini, 2012).
The model of Forsythe et al. (1995) was used to determine day

length (including twilight) and the time of dawn and dusk based
on the day of the year (DayofYear) and latitude.
Daylight = sin [(T 0.25) 2] 0.5
DayTwlength
24
(1)
where Daylight assumes a positive value when it is light and a negative value when it is dark. T is time of day (unit: days). DayTwlength
(hrs.) is the length of the day including twilight and calculated from
the day of the year and latitude as:
DayTwlength = 24
24
acos (DayTwlengthP2)
(2)
where,
DayTwlengthP2 =
sin 6/180 + sin Latitude /180 sin (DaylengthP1)

cos (Latitude /180) cos (DaylengthP1)
(3)
and DaylengthP1, the daylength excluding Twilight hours for the

lactation module in Molly,
DaylengthP1 = asin 0.39795 cos 0.2163108 + 2 atan

tan [0.0086 (DayofYear 186)]

0.9671396
(4)
14
These equations do not perfectly reect the length of day at latitudes greater than 60 . Thus, a set of conditional statements are
used to cap daylength to a value of not greater than 24 h (Appendix
A.2.1).
of ruminal ll and plasma ghrelin in grazing dairy cows. Ruminal

ammonia has been related to the control of herbage intake and its
diurnal pattern (Vuuren, 1993; Chapman et al., 2007). Conrad et al.
(1977) also reported reductions in intake rate and meal duration
as ruminal ammonia increased. High ruminal ammonia has been
negatively related to neuropeptide-Y and ghrelin secretion (Miner,
1992). AHor is a proxy for insulin (Hanigan et al., 2009), which has
a negative feedback effect on intake, inhibiting feeding motivation
and decreasing meal size (Asarian and Geary, 2006; Roche et al.,
2008a), and indirectly related (negatively) to ghrelin secretion in
dairy cows (Roche et al., 2008a).
KFdRat (from Eq. (7)) includes a lag function based on the equation
of Roseler et al. (1997) to represent the reduction of intake in early
lactation functionally. KFdRat is calculated as follows:
2.2.2. Hunger hormone

As a meal progresses, hunger signals (opposite of satiety)
derived from the gut and absorbed nutrient supply decrease, which
contributes to cessation of a meal and changes in ingestive behaviors (Dufort and Wright, 1962; Teitelbaum, 1966; Allen et al., 2005;
Gregorini et al., 2009c). Many of these signals communicate with
the brain via hormonal circulation in the blood and via peripheral
nerves (Morton et al., 2006; Roche et al., 2008b), and are integrated
with signals coming from the animal (e.g. adipose tissue (Faverdin
et al., 1999; Geary, 2004)) and external stimuli (Smith, 1996) to
determine the level of feeding motivation (Fig. 1) (Rhind et al., 2002;
Gil-Campos et al., 2006; Gregorini, 2011).
Hunger is represented in the model by a hormone denoted as
IHor (hunger hormone), which controls the onset and cessation of
meals. Although IHor represents an aggregated representation of
stimuli, it is roughly patterned after ghrelin. The orexogenic properties of ghrelin and its relationship with the onset, cessation and
intra meal eating behavior are well documented (Miner, 1992; GilCampos et al., 2006; Roche et al., 2008b; Gregorini, 2011). Increased
IHor triggers a meal, while decreased IHor causes eating cessation. IHor also modulates ingestive behavior within each meal as
described in Section 2.5. IHor at any point in time is determined
by numerical integration of the differential equation describing
synthesis and degradation:
d (IHor)
= IHorSyn IHorDeg
dt
kFdRat = 1 +
(6)
where iIHor represents the reference concentration of IHor; set to

1 as for other hormones in Molly.
The differential equation describing IHor synthesis (IHorSyn) is:
IHorSyn =
1 + CVFA /kVFA kFdRat
xVFA
+ AHor/kaHor kFdRat
xaHor
MamCellsPart
1000
(8)
where, BCSTarget , is the representation of the target adipose tissue

weight defended by the animal depending on the stage of lactation.
BCS BCSTarget MamCellsPart, and LagDMI are body condition score
target, actual BCS, number of milk secretory cells in the udder at
parturition and the function representing the reduction of intake
in early lactation, respectively.
Body condition score is used to represent changes in adiposity and thereby leptin secretion (Vega et al., 2013; Roche et al.,
2009), and MamCellsParts represents genetic merit (potential for
milk production) of the cow (Hanigan et al., 2008), respectively.
Adipose tissue dynamics and BCS are dynamically represented in
Molly (Section 2.1). Leptin acts to maintain adipose tissue reserves
(Roche et al., 2008b), and thus leptin concentrations and BCS are
positively correlated (Vega et al., 2013; Delavaud et al., 2000; Len
et al., 2004). Leptin also controls feed intake (Vega et al., 2013;
Faverdin et al., 1999; Delavaud et al., 2000) because reduced leptin
secretion stimulates feeding. The effect of leptin is thought to be
mediated by glucose and ketone bodies (Delavaud et al., 2000), as
(5)
(dIHor) + iIHor
kBCS +
kMamCells + LagDMI xLag 1
IHor =
BCSTarget
1
BCS
VmIHorSyn
+ RumDM/kRumDM kFdRat
xRumDM
+ Am/AmCor/kAm kFdRat
xAm
(7)
where, VmIHorSyn , CVFA , RumDM, and Am represent, respectively, the

maximum velocity of IHor synthesis, concentration of volatile fatty
acids (VFA), DM load (kg, dry matter basis) in the rumen and ammonia (mol/L). AmCor is a correction factor for Am. AHor represents
blood anabolic hormone, while kFdRat is a function adjusting for
changes in adiposity and genetic merit for milk production (see
(Hanigan et al., 2009, 2013) for further details on these variables).
The rationale for the various factors affecting IHorSyn is as follows. Ruminal volatile fatty acids (VFA) are major satiety signals. Of
the major VFA produced in the rumen, acetate and propionate are
thought to produce the strongest satiety signals (Faverdin, 1999).
While propionate seems to be the strongest signal (Farningham
and Whyte, 1993) the concentration and rate of total VFA production in the rumen are considered to be the main satiety signals
(Adams and Forbes, 1981; Illius and Jessop, 1996); they are negatively related to ghrelin secretion (Roche et al., 2007; Gregorini,
2011). Since the work of Cannon and Washburn (1912), for example, the ll effect is also considered a major factor in determining
the amount and rate of feed consumed at each meal, meal ingestive behavior (Baumont et al., 1990; Gregorini et al., 2007a,b, 2009c)
and the diurnal arrangement of meals (Gregorini, 2012). Gregorini
et al. (2009c) reported negative relationships between the level
well as the orexigenic neuropeptide-Y (Schwartz et al., 1996; GilCampos et al., 2006). There is an interaction between leptin and the
neuropeptide-Y (Gil-Campos et al., 2006) that explains the negative relationship between leptin and ghrelin secretion (Stylianou
et al., 2007). Leptin does not dictate the onset or cessation of a
meal, it modulates feeding motivation (Houseknecht et al., 1998).
Roche et al. (2006) observed that genetic selection for milk production increased plasma ghrelin concentrations and associated
herbage intake levels in dairy cows. This mechanism is represented
by MamCellsPart, which represents secretory cell numbers and differentiates udder metabolic capacity between cows, enabling the
model to represent genetic merit for milk production.
A reduction of intake in early lactation (LagDMI) is generally
observed in dairy cows. This relates to reductions in gut capacity
and is potentiated by adipose tissue mobilization (utilization) and
status (Faverdin et al., 1999). The lag function described by Roseler
et al. (1997) captures this phenomenon.
IHor degradation is calculated as a mass action function of IHor,
kIHor being a constant for adjustments.
IHorDeg = kIHor IHor
(9)
Having dened IHor, trigger points for starting (StartIHor) and

ending (StopIHor) a meal were determined from iIHor by dening
a range (IHorRange),
StopIHor = iIHor
IHorRange
2
StartIHor = iIHor +
IHorRange
2
(10)
A meal will start (IHor > StartIHor) and stop (IHor < StopIHor) subject to additional constraints executed as a series of logical tests
(Appendix A.2.2). These include: (1) a primary meal cannot start
when Daylight < 0; (2) once started, a meal can continue past dusk;
(3) a small meal can occur after dusk, and (4) a meal cannot occur
when the cow is out of the paddock and/or being milked. Because
StopIHor and StartIHor are not equal, the time between meals and
the length of a meal are primarily determined by the magnitude
of the range between the triggers (IHorRange), creating an intermittent grazing pattern. Because meals occurring during the late
afternoon and early evening are generally longer and more intensive compared to previous meals of the day (Gibb et al., 1998;
Taweel et al., 2004; Gregorini, 2012), StopIHor was adjusted downwards 3 h before the end of the day, a period labeled Latefeeding,
and a logical statement was used to reduce the StopIHor during that
period (Appendix). Latefeeding was determined from DayTwlength;
Latefeeding = sin [(T 0.125) 2] 3 0.5
DayTwlenght
(11)
24
2.3. Diurnal pattern of rumination

Similar to the model of Sauvant et al. (1996), rumination occurs
as a response to the size of the large particle pool in the rumen.
The fraction of large particles swallowed is determined and used as
an input to the pool (Fig. 2). During inter-meal periods, rumination
occurs if the pool of large particles (LP) is greater than the minimum
LP pool size (MinLPRumntn). Rumination continues until the LP pool
falls to MinLPRumntn or another meal begins. When ruminating,
the rate of particle breakdown to small particles is represented as a
mass action function (in Molly) governed by the rate constant kLPSP .
When MINDY is not grazing or ruminating, MINDY is considered to
be resting (Appendix, A.2.4.). Daily rumination time is calculated
by numerical integration.
Rumination is important, not only because of the physical
breakdown of ingested feed particles, but also because it affects
salivation. Saliva supplies between 70 and 90% of the uid and
buffering capacity entering the rumen and is the major determinant of liquid outow rate from the rumen (Cassida and Stokes,
1986). Observations from Maekawa et al. (2002) and Cassida and
Stokes (1986) were used to set the previously encoded salivation
rates associated with eating, ruminating, and resting (225, 205 and
114 mL/min, respectively) in Molly code.
sward canopy is relatively homogeneous (horizontal plane) and

the area offered and time allocated to graze are restricted, cattle
graze down available herbage in successive strata (Ungar and Ravid,
1999). Even in relatively homogeneous swards, canopies present
vertical chemical and morphological heterogeneity, which changes
throughout the day (Delagarde et al., 2000; Gregorini, 2012). This
creates different spatio-temporal arrangements of availability and
accessibility of herbage mass and nutrients. Therefore, as cattle progressively graze down a pasture, they dene and then encounter
successive and dynamically changing grazing strata with respect
to feeding value.
2.4.1. Characteristics of the sward canopy, grazing strata and
chemical composition of herbage
The sward canopy is modeled as a set of superimposed grazing strata (GS) as in Galli et al. (1999) and Baumont et al. (2004).
The sward canopy depth (m) determines the vertical dimension of
the sward canopy available to be grazed, which is determined by
the initial extended tiller height (ETHini ), an input to the model.
The number of accessible GS (Nstrata) within the sward canopy
available to be grazed is calculated as follows:
Nstrata = 1 +

MeanLM =

[Log (ETHlim ) Log (ETHini )]

Log (1 DD)
(12)
where, ETHini (m) and ETHlim (m) are the initial extended tiller
height and the sward canopy barrier, respectively. The barrier is the
sward canopy height under which cows are not allowed to (determined by management) or able to graze (determined by sward
canopy structure; i.e. stem/pseudostem physical barrier). ETHlim is
an input to the model. DD, a constant proportion of ETHini , represents bite depth for each GS (Wade et al., 1989), thus dening the
depth (m) and ETH (m) of each GSi in the sward canopy. The latter
is calculated as:
ETHi = ETHini (1DD)(i1)
(13)
For the purpose of calculations of vertical distribution of herbage

mass and nutrients within the sward canopy, the median point
height of each GS (Hi , m) is calculated as:
Hi =
[ETHi + ETHi1 ]
2
(14)
Herbage biomass (kg DM/m2 , input to the model) varies through

the vertical plane of the sward canopy. Such a distribution is
described by a morphological analysis (mass, position and morphological components) similarly to Baumont et al. (2004), utilizing a
linear mass index (LMIi ) that describes the vertical distribution of
herbage mass within the sward canopy.

1
LMIi =
MeanLM
2.4. Grazing environment and behavior

Grazing ruminants prefer to graze swards and generally select
patches that enable them to maintain or increase herbage intake
(Demment et al., 1993; Illius et al., 1999). Therefore, when the
15
LM +

(SM LM)
1 + MeanSwardHeight/Hi
eLMI
(15)
where, LM and SM are the masses of lamina and sheath plus stem
(kg DM/m) respectively. MeanSwardHeight is half of ETHini and eLMI
is a parameter that changes the shape of the curve and so the
distribution of herbage mass in the vertical plane, allowing the
representation of different vertical distributions. MeanLM is the
mean linear mass (kg DM/m) of a grass tiller plus stem (or erect
legume) calculated from the integration of the MichaelisMenten
curve between ETHini and MeanSwardHeight.
[(ETHini + MeanSwardHeight) LM] + MeanSwardHeight eLMI (LM SM) / (ETHini + MeanSwardHeight)

MeanSwardHeight LM + (MeanSwardHeight eLMI (LM SM) /MeanSwardHeight)
ETHini
(16)
16
While herbage mass is an input to the model, mean bulk density

of the sward canopy (MBDsward , kg/m3 ) and each GS (MBDi , kg/m3 )
are calculated as:
MBDsward =
HM
ETHini
MBDi = MBDsward LMIi
(17)
(18)
Temporal changes in the nutrient composition of herbage mass

are represented using a set of polynomial equations to predict
changes in neutral and acid detergent bers, crude protein, soluble crude protein, and an estimate of the rumen undegradable acid
detergent ber concentrations continuously during daylight hours
(Appendix A.3). The equations are of the form:
FAdjustment = Cons tan t + a T 3 + b T 2 + c T
(19)
where FAdjustment is the adjustment factor to the herbage chemical composition, T represents time of day (days) calculated using a
MOD function; a, b and c are coefcients derived for each nutrient
from the data of Gregorini et al. (2008c, 2009d). FAdjustment was centered on a value of 1. As composition data were only available for
daylight hours, the adjustment factors were calculated as a linear
interpolation of the value at sunset and the value at sunrise during
dark hours. Thus, the adjustments cycled each day. Fiber and protein concentrations (g/kg DM) were then calculated continuously
from the herbage chemical composition inputs as:
NutrientAdjusted = Nutrientinput FAdjustment
(20)
Soluble carbohydrates are calculated by difference after specication of all of the other nutrients in the herbage. Thus, net changes
in the sum of temporally adjusted and explicitly dened nutrients
were inversely proportional to the soluble carbohydrate content
of herbage. Changes in biomechanical (e.g. toughness) features of
herbage affecting ingestive behavior are represented by the neutral detergent ber content. Neutral detergent ber content varies
in the vertical plane of the sward canopy and by time of day. These
spatio-temporal changes were represented using the equation of
Baumont et al. (2004) with temporal adjustment using FAdjustment
for neutral detergent ber (FAdjustmentNDF ):
NDFi = (1.15 7.33e 4 Hi) NDFadjusted
(21)
where, FAdjustmentNDF is dened by Eq. (19). The NDF content is used

in the model (see Molly code) to calculate nutrient inputs like hemicellulose, cellulose, and soluble carbohydrates. Thus, as NDF varies
in time and space (by strata), reciprocal changes in herbage soluble carbohydrates, hemicellulose and cellulose occur across the
vertical plane of the sward canopy.
2.4.2. Potential and actual herbage intake rate at each grazing
stratum
The short-term herbage intake rate (kg DM/unit of time) of a
grazing animal can be represented by the product of two variables
bite mass (BM, kg/bite) and bite rate (BR, bites/min). In the model,
as in Baumont et al. (2004), the potential herbage intake rate at
each GS, (PIRi , kg DM/min), is calculated as:
PIRi =
BMi
TBi
(22)
where, is BMi given by:

BMi = BDi BAi MBDi
where BDi and BAi are bite depth (m) and area (m2 ) and MBDi
mean bulk density (kg/m3 ). BDi and BAi are calculated as:
BDi = ETHi DD
(23)
is the
(24)

BAi =
2 DA2
1 + MeanSwardHeight/Hi

exp [0.3 (MBDi 1)] (25)
where DA (m) is the animals dental arcade width, which is calculated based on a modication of the Illius and Gordon (1987)
allometric relationship to avoid the effect of gut ll, adipose tissue
and gravid uterus on animal body weight (BW, kg):
DA = 8.6 NonFatNonUterEmptyBW 0.36
(26)
Because BA is dependent on DA (Illius and Gordon, 1987) and

sward surface height, the asymptote of this relationship was set to
twice the square of the DA (Gordon et al., 1996) and affected by
MeanSwardHeight and Hi as in Baumont et al. (2004). The negative
exponential function represents the effect of sward density on BA,
which has a neutral effect for a MBDi of 1 kg herbage dry matter per
m3 (Laca et al., 1992).
The potential time per bite (TBi , min) is the sum of prehension (PT, min) and potential ingestive chewing times (PCHT, min)
for each GS, where the latter is a function of the herbage neutral
detergent ber content and BM:
TBi = PT + PCHT
(27)
PCHTi = (0.25 NDFi 12.5) BMi
(28)
where, PT is based on Illius and Gordon (1987) using empty BW

minus fat and uterine weight (NonFatNonUterEmptyBW) so that
changes in gut ll, BCS, and pregnancy state do not affect the estimate:
PT = 0.35 NonFatNonUterEmptyBW 0.125
(29)
Ingestive chewing is a variable function of the mass and brosity of herbage and hunger (Prez-Barbera and Gordon, 1998).
Grazing cattle increase TB as BM and ber content of harvested
herbage increase, but reduce it as hunger increases as a compensatory mechanism to increase herbage intake rate (Greenwood and
Demment, 1988; Gregorini, 2011). Therefore, from TBi and IHor, the
actual chewing time (ACHT, min) required for each bite taken is
calculated as:
ACHTi = (TBi PT ) ChewFactor
(30)
where, ChewFactor is a variable representing the effect of feeding

motivation on ingestive chewing [See (Prez-Barbera and Gordon,
1998; Gregorini, 2011)] and calculated as:
ChewFactor = LowChewingMot [(LowChewingMot
HighChewingMot) exp[kChewfactor GIHor] ]
(31)
where LowChewingMot and HighChewingMot are constants with

values of 0.15 and 1, respectively, and kChewfactor is a parameter that
changes the shape of the curve and so the time invested in ingestive
chewing at different levels of hunger as a function of external factors such as available herbage mass (AHM), time of day, and GIHor
(See Section 2.4.3).
Therefore, the actual herbage intake rate (kg/min) for a particular GSi at a given time of day (t) is calculated as:
ActualIRi =
BMi
PT + ACHT
(32)
2.4.3. Actual motivation to graze

Modulation of hunger (IHor) by indirect controls (external
stimuli, incentives) represents (to some extent) the animals perception of the feeding environment, which can change ingestive
behavior. Hunger and, thereby, ingestive actions are modulated
by time of day, herbage chemical composition, available herbage,
light intensity and anti-predator strategies (Woods and Strubbe,
17
1994; Strubbe and Woods, 2004; Gregorini, 2012). Therefore, actual

motivation to graze is calculated as:
GIHor = IHor AHM TimeOfDayEffect
(33)
where, AHM is the available herbage mass modulator factor at a

particular T. AHM is calculated as:
AHM =
(HMtotavail + HMunavail )
iHMtotal
HMtotavail =
(GSAi i MBDi i) BDi
(34)
(35)
HMunavail = iHMtotal iHMtotavail
(36)
iHMtotal = HM TA
(37)
iHMtotavail = TA MBDsward (ETHini ETHlim )
(38)
where, HMtotalavail is the sum of herbage mass (kg) remaining in

each GSi . HMunavail is that herbage below ETHlim . iHMtotal is the total
pre-grazing herbage mass (kg/m2 ). TA is total grazing area allocated
per cow (m2 ). iHMtotavail is the AHM at the time pasture is allocated
(pre-grazing herbage mass).
TimeOfDayEffect (unitless) was patterned after the natural diurnal grazing patterns of ruminants, using data (patterns of ingestive
behavior) from set stocked dairy cows (Gibb et al., 1998; Taweel
et al., 2004), sheep (Orr et al., 1997) and beef heifers (Gregorini et al.,
2007a). It stimulates the motivation to graze as dusk approaches
(Gregorini, 2012) and reduce it overnight. TimeOfDayEffect calculus
is presented in the Appendix (A.2.6).
2.4.4. Grazing strata, foraging decisions and actual herbage
intake dynamics
2.4.4.1. Dynamics of grazing strata area availability and accessibility.
As in Baumont et al. (2004), a GS is considered relatively homogeneous and thus can be represented as cluster of contiguous bites
(Fig. 3a). Therefore, if iHMtotavail occupies the TA, each grazable
stratum area (GSA, m2 ) initially equals the sum of the potential BA
in the GS. As the pasture is grazed down, the next lower GS will progressively become accessible. Therefore, GSA accessibility for each
GS changes as grazing occurs.
dGSAi
= SGRi CRi
dt
(39)
where, SGRi and CRi are the rates of change in the area (m2 ) of
each GS due to sward canopy growth (m2 /min) and consumption,
respectively (Also see Fig. 3a). SGRi and CRi are calculated similarly
to Baumont et al. (2004).
[(TAGSAi ) HGR] dGSAi
= SGRi CRi
SGRi =
BDi
dt
(40)
CRi = BRi BAi
(41)
where GSAi (m2 ) is the ungrazed and accessible area for GSi
(derived
by numerical integration of Eq. (39)); HGR is the herbage growth
rate (m/min); and BRi is bite rate (bites/min) for GSi (see Section
2.4.5).
2.4.4.2. Foraging decisions. As successive GS become accessible, the
animal can either (1) completely graze the upper GS before starting
to graze the next lower GS, or (2) graze completely down the sward
canopy at each feeding station (FS, Section 2.4.6) before moving to
the next FS. The former is consistent with an attempt by the animal
to maximize intake rate given that BM successively becomes less
in each lower GS (Demment et al., 1993; Illius et al., 1999). Under
rotation grazing management, cattle graze by stratum (Wade et al.,
Fig. 3. a) A schematic representation of the bite features and grazing strata during
grazing; b) a schematic representation of the harvesting step length calculus.
(Adapted from Galli et al. (1999) and Baumont et al. (2004)).
1989). They start consuming the next lower GS as the area of the
upper stratum shrinks. In MINDY, it was assumed that the animal
would only graze from the upper two available GS, and the preference for each strata (PREFi and PREFi 1 ) was calculated from the
proportion of the upper GSA remaining using a MichaelisMenten
function to achieve a continuous shift in preference from the upper
to the lower GS as the GSA of the upper stratum declined. A series of
logical statements (Appendix A.2.5.) was used to shift grazing down
to the next pair of GS (i.e. GSi 1 and GSi 2 ) when GSAi reached a
negligible area (minimumGSA).
2.4.5. Real herbage and nutrient intake and bite rate
Herbage intake rate (FdRat, kg/min) is a function of ActualIR and
PREF for each of the pair of available GS:
FdRat = (PREFCurrentStratum ActualIRCurrentStratum + PREFCurrentStratum1
ActualIRCurrentStratum1 )
(42)
where PREFCurrentStratum and PREFCurrentStratum 1 are the relative

preferences for the upper and next lower GS, respectively.
Therefore, the rate of each particular nutrient intake is calculated as:
FdRatNutrient = FdRatCurrentStratum NutrientCurrentStratum + FdRatCurrentStratum1
NutrientCurrentStratum1
(43)
where, NutrientCurrentStratum is calculated as described in Section

2.4.1. Eqs. (1921).
Daily herbage DM intake (HDMI, kg/d) and particular nutrient
intake (kg/d) are determined by numerical integration, for example:
t
HDMI =
FdRat
0
(44)
18
Having determined PREF, ActualIR, and BM, bite rate (BR,

bites/min) can is calculated as:
BR =
PREF
CurrentStratum ActualIRCurrentStratum
BMCurrentStratum
PREF
CurrentStratum1
ActualIRCurrentStratum1

BMCurrentStratum1
HSL = CowHeight tan 30
FSR =
(46)
(47)
HSL
The momentary average speed while harvesting (MSH, m/min)

is derived as:
MSH = FSR HSL
(48)
and the distance walked while harvesting (DWH, m)
t
DWH =
MSH
(49)
Searching time (STI, days) and distance (SDI, m) covered while

searching are calculated assuming that maximum searching velocity (MVelsearching , m/min) also changes in response to grazing
motivation as shown by Gregorini et al. (2007a,b). Considering that
the grazing process is equal to harvesting plus searching, searching
time is calculated only when the cow is grazing but is not harvesting
herbage:
pSTI = 1 [BR (PT + ACHT )]
(50)
where, pSTI momentary proportion of time searching (unitless), and

STI calculated as:
t
STI =
pSTI
(51)
Then, momentary average searching speed (SSpeedS, m/min)
SSpeedS = pSTI
MVelsearching
GIHor
SSpeedS
(53)
(45)
where, CowHeight (m) is the shoulder height of the cow.

Assuming random searching for FS while grazing, a maximum
walking speed while harvesting (MVeleating , m/min) (Ungar and
Noy-Meir, 1988; Galli et al., 1999) and that such a velocity slows
down as grazing motivation declines (Gregorini, 2011; Gregorini
et al., 2011), the momentary rate of FS (FSR, FS/min) is calculated
as:
MVeleating GIHor
t
SDI =
2.4.6. Walking and searching while grazing

Grazing includes a series of animal decisions at different scales
(Senft et al., 1987). Once the animal has made the decision to graze,
it selects an area to place the bites, exploits it and then abandons
it. Grazing encompassed two main actions, harvesting (prehension,
ingestive chewing and swallowing) and searching. A feeding station
(FS) is the area of pasture a grazing animal can reach from a given
position without taking a step. The number of FS would then be
equal the number of steps taken while the cow is head down harvesting (Ruyle and Dwyer, 1985; Rook et al., 2004; Wade et al.,
2006; Gregorini et al., 2011). The area of a FS is calculated as a semicircle, where the step length while harvesting (HSL, m) is used as the
radius (m). Based on the works of Ruyle and Dwyer (1985) and Rook
et al. (2004) and assuming a 30 angle for the neck when the cow
is head down harvesting, the HSL is trigonometrically calculated as
(Fig. 3b):
and SDI calculated as:
(52)
Therefore, the searching step rate (SSR, searching steps/min) and

the total number of steps while searching (TSS) are calculated as:
SSR =
SSpeedS
(HSL 1.4)

TSS =
x
y
(54)
SSR
yx
(55)
where, 1.4 is a scalar to calculate searching step length (m) which

is 40% longer than HSL (Gregorini et al., 2007a,b).
During lactation, in addition to HTD, cows walk from the pasture
to the milking parlor. This extra walking (RacesDistance, m) depends
on milking frequency and distance to the parlor. Therefore, it is
possible to calculate the total distance walked during a particular
day (DailyWalking, m).
DailyWalking = DWH + SDI + RacesDistance milking frequency
(56)
3. Model parameterization
Due to the complex interactions among the sub-models, multidimensional sensitivity analyses were run to identify patterns of
change in HDMI, grazing time, rumination time, resting time, number of meals per day, and meal length in response to variables
affecting the diurnal pattern of IHor. Model parameters were initially set to achieve the approximate number and length of meals
per day. Variables were perturbed in both a positive and a negative direction from their assumed values by 20% in two steps. Then
model outputs were regressed on the change in the variables estimates to estimate the model sensitivity (Appendix A.4, Table A.3).
Herbage dry matter intake and daily grazing time responded to
most of the variables tested as expected. They were particularly
well correlated with changes in parameters controlling IHor synthesis and degradation and the minimum interval between meals at
night. Rumination time was sensitive to IHor synthesis and degradation, the multiplier to expand the IHor range during dusk, the
rate of reduction of large particles to small particles in the rumen,
and the pool of DM in the rumen. Resting time mirrored rumination time. Meals per day was most sensitive to factors controlling
IHor synthesis and degradation, like anabolic hormone, pool of DM
and VFA concentration in the rumen. Intake per meal was not sensitive to any single parameter but was moderately sensitive to
IHor synthesis, reduction of large particles to small particles in the
rumen, and VFA and ammonia concentration in the rumen. The
length of meals was not sensitive to any of the parameters but was
moderately sensitive to IHor synthesis and to VFA and ammonia
concentration in the rumen. This test provided a better understanding of how each of the variables affected model outputs, thereby
identifying which parameters determined total intake and patterns
of intake within a day.
Parameter estimation was conducted by tting the model to the
data of two experiments conducted with grazing dairy cows under
set stocking grazing management, Gibb et al. (1998) and Taweel
et al. (2004), since under this management, domestic ruminants
show a natural grazing pattern Gregorini (2012). The experiment
of Gibb et al. (1998) was conducted in summer and used Friesians
cows in mid-lactation (160 days in milk) with an approximate
liveweight of 470 kg, grazing a L. perenne dominated sward of
57 cm surface height and an approximate mass of 2200 kg DM/ha.
Cows were milked twice a day at 5 am and 2:30 pm. The experiment of Taweel et al. (2004) was conducted in the spring and
used Holstein cows in late lactation (310 days in milk) with an
approximate liveweight of 600 kg, grazing a L. perenne dominated
sward of approximately 1015 cm surface height and a mass of
1700 kg DM/ha. Cows were milked twice a day at 6:30 am and 6 pm.
The model inputs were congured to reproduce the above sward
characteristics, time of year, latitude, and milking schedule. The
observed discrete patterns of grazing activity were reconstructed
using a time scale of minutes and smoothed to create a continuous
variable using the following equation. This was required for parameter estimation as the available optimizers in ACSLX are not able to
handle discrete data.
dGrazingSmotothed =
(SmoothingCons tan t time step + Grazing)

(1 + SmoothingCons tan t time step)
t
GrazingSmoothed =
dGrazingSmoothed
0
where GrazingSmoothed is a moving average of grazing activity with

a ratio of 10 to 1 for the previous moving average, SmoothingConstant is 10 and grazing is the discrete meal patterns shown by the
data.
The same equation form was used in the model to convert
the predicted discrete meal patterns into a continuous variable
which was then compared to the function values derived from
the observed data. Fig. 4 shows a plot of smoothed meal pattern
for observed and predicted values. Parameters for estimation were
chosen based on the sensitivity analyses and estimated using the
NelderMead algorithm available in ACSLX.
Model parameter estimates and an analysis of residual errors are
presented in Table 1. In all cases, the model parameters were well
dened by the data as evidenced by the STD of the parameter estimates being less than 1% of the respective estimates. Synthesis of
IHor was most sensitive to changes in ruminal VFA concentrations
as evidenced by the sensitivity exponent of 10.35. Sensitivity of the
equation to changes in AHor, ruminal ammonia, and ruminal DM ll
were more modest assuming sensitivity exponents near the initial
values of 1. The evening and night-time range between StopIHor
and StartIHor was found to expand to 1.24 times the range during
the daytime. This helped trigger initiation of a late afternoon, early
evening meal and expanded the length of such a meal to ensure
adequate ruminal mass through the night. The length of meals initiated after dark was limited to 64 min. With the derived settings, the
model had 36% error in predicting the timing and length of the various meals throughout the day as reported in Gibb et al. (1998) and
19
Taweel et al. (2004). The model did not display mean bias (0.35%
of the MSPE) and slope bias was minor (20% of the MSPE). This
suggests that most of the variance was due to random biological
variation (79% of the MSPE).
4. Illustration, application and discussion

The present work focused on formulating and parameterizing
the model with an initial validation conducted under contrasting
scenarios. This validation was performed by assessing its ability
to simulate realistic diurnal grazing patterns and herbage intakes
for dairy cows of two distinct genotypes, old (1970s origin)
and modern (1990s origin) New Zealand HolsteinFriesian in
mid-lactation (late spring) subjected to two contrasting grazing
managements: (1) set stocking and (2) rotational grazing (24 h pasture allocation) on a L. perenne dominated sward. Stocking rate was
the equal for both management scenarios; while grazing pressure
(stock density per unit of area over a 24 h period) was lower for
set stocking. In rotational grazing the area allocated per cow per
day was 100 m2 , while the area available to graze over 24 h in set
stocking was 2000 m2 (equivalent to a rotation length of 20 days).
The parameters and inputs used to describe cows and swards are
presented in Table 2. The results of the simulation are presented in
Table 3 and Fig. 5 and discussed in the following sections.
4.1. Daily herbage intake and hunger level

The results of the simulation show a marked effect of animal
genotype on daily herbage intake. The predicted level of intake,
and the magnitude of its difference between cow genotype, agree
with the empirical data of Macdonald et al. (2008) collected under
New Zealand grazing conditions and management. Macdonald et al.
(2008) reported intakes of 13.9 and 13.7 and 16.8 and 15.7 kg for the
old and modern cows at 90 and 240 days in milk, respectively. Previous empirical modeling (Gregorini et al., 2009b) for these two cow
genotypes indicated that modern cows of NZ origin have greater
feeding drive (hunger) in comparison with the older cows type
which is consistent with Roche et al. (2006) who reported greater
level of plasma ghrelin for modern cows (301 vs. 241 pg/mL, respectively). MINDY predicted not only a different diurnal pattern of the
hunger hormone (IHor ghrelin) across cow genotypes, especially
under rotational grazing management (Fig. 5), but also different
daily means (Table 3). Increments in intake come with genetic
improvements for milk production (Linnane et al., 2004; Kolver
et al., 2005). Although detailed statistical validation is required, the
present example suggests that MINDY can predict realistic levels of
Table 1
Model parameter estimates derived by tting to the data of Gibb et al. (1998) and Taweel et al. (2004), and residual error analyses.
Parameter1
Final
STD of estimates
Residual error analyses
VmIHorSyn
kIHor
NightMealInter
NightMealTime
NightMult5
kLPSP
kaHor
kAm
kRumDM
kVFA
xaHor
xAm
xRumDM
xVFA
73.26
6.01
0.10
0.04
1.24
2.78
1.53
2.99
3.38
0.11
0.85
1.16
1.02
10.35
0.0027
0.009
0.00001
0.000003
0.003
0.00081
0.00023
0.00013
0.00047
0.000033
0.00036
0.00025
0.00016
0.0056
RMSPE, % of mean
Mean bias, % of MSPE
Slope bias, % of MSPE
Random error, % of MSPE
N
Mean of observed
Mean of predicted
RSMPE, square root of mean prediction error; STD, standard deviation; MSPE, mean square prediction error.
1
Parameters are described in Table A.1.
36.18
0.35
20.71
78.92
5264
0.46
0.47
20
Fig. 4. Plot of smoothed observed values from Gibb et al. (1998) and Taweel et al. (2004), overlaid with the smoothed predicted values for grazing activity (i.e. meal throughout
the day).
Table 2
Input parameters used to represent cow genotype and sward features for simulations of cows with historic or current genetic potential subjected to set stocked or rotational
grazed pastures.
Sward characteristicsa
Parameter
Herbage mass (kg DM/ha)

Extended tiller height (cm)
eLMI (unitless)
SM (g/cm)
Area available to graze over 24 h (m2 )
Herbage growth rate (cm/d)
Crude protein (g/kg herbage DM)
Lipids (g/kg herbage DM)
Non-structural carbohydrates (g/kg herbage DM)
Neutral detergent ber (g/kg herbage DM)
Acid detergent ber (g/kg herbage DM)
Lignin (g/kg herbage DM)
Ash (g/kg herbage DM)
Set stocking
Rotational grazinga
2300
15
0.00001
0.001
2000
0.6
178
40
60
385
206
16
100
3000
30
1
0.0015
100
0.6
178
40
60
385
206
16
100
Cow genotype
Mature weight (kg)

Height (from ground to shoulders, m)
Liveweight (kg)
MamCellPart (unitless)
Body condition score (points, 15 scale)
Days in milk (days)
Milking times (hours)
a
Old
Modern
500
1.4
440
710
3.25
180
5 am, 3 pm
550
1.6
470
860
3.0
180
5 am, 3 pm
Strips of new pasture were allocated every 24 h right after morning milking (5:30 am). Chemical composition of the grass at noon (12:00 pm).
Table 3
Effect of cow genotype and grazing method on model predictions of variables associated with hunger, dry matter intake, grazing pattern, and distance walked.
Cow genotype
Old
Modern
Grazing method
Set stocking
Rotational grazing
Set stocking
Rotational grazing
Output variable
IHor (unitless)
Herbage intake (kg DM/d)
Grazing time (min/d)
Harvesting time (min/d)
Searching time (min/d)
Number of meals per day
Average meal length (min)
Average meal intake (kg DM)
Rumination time (min/d)
Idling time (min/d)
Distance walked while eating (m/d)
Distance walked while searching (m/d)
0.91
15.2
492
487
5
5
102
3.04
442
505
2444
30
0.99
15.3
485
297
187
5
59
3.01
471
483
3192
1044
0.95
17.8
544
537
7
5
115
3.55
441
455
2455
33
1.43
16.6
560
351
208
5
75
3.32
435
446
3357
956
21
Fig. 5. Diurnal pattern of grazing behavior, rumen function and anabolic and hunger hormones of two genotype of cows [Old (a) and Modern (b)], under two grazing methods
[set stocking and rotational] as simulated by MINDY model. (Solid lines belong to y axis and dotted lines belong to the z axis).
herbage intake (Table 3), and thus be used to evaluate the response
in herbage intake of cows with different genetic merit.
Grazing management also altered the level of intake. Herbage
intake was lower for the modern cow under rotational grazing
as a result of herbage depletion, which constrains compensatory

ingestive behavior and thereby herbage intake rate, as shown in
Fig. 5b and supported by Dillon (2007). Relatively small differences
in herbage intake were predicted between grazing methods for
22
the old cow type. This prediction matches previous observations

(McMeekan and Walshe, 1963; Bluett and Macdonald, 2001) of a
small, or no, difference in herbage intake between these grazing
methods when availability of herbage is not limiting and feeding
drive is relatively low. This premise is reinforced by the predicted
daily means of IHor (Table 3), in other words, hunger.
4.2. Daily and diurnal patterns of ingestive behavioral activities
4.2.1. Grazing
Predicted daily grazing times by MINDY fell within the ranges
(481608 min/d) reported in a meta-analysis by Prez-Prieto and
Delagarde (2012) for grazing dairy cows with similar characteristic
and feeding in similar grazing environments to the ones simulated
for the present illustration. More specically, MINDY predicted different grazing times, along with different harvesting and searching
time budgets, depending on cow genotype and grazing management (Table 3). These predictions follow the pattern observed by
Rossi et al. (2004, 2005), who reported different daily grazing times
for the old and modern cow genotypes during early lactation, 489
and 509 min/d, respectively. These considerations indicate realistic prediction of daily grazing time by MINDY and its potential to
represent grazing time as affected by cow genotype.
MINDY predicted consistently different mean meal length and
herbage intake per meal for cows of different type subjected to different grazing management (Table 3). When combined with Fig. 5,
the data indicate distinct grazing, herbage and nutrient intake patterns. Modern cows demand more nutrients, leading to longer,
more intense meals, as previously suggested by Roche et al. (2006)
and Linnane et al. (2004). Rotational grazing reduced the length
and level of intake of the average meal. These two meal features,
plus within meal ingestive behaviors (bite rate, chewing time, eating and searching steps rates) and their diurnal pattern follow the
diurnal grazing pattern described by Gregorini (2012). Set stocked
ruminants show the longest and most intensive meal during the
afternoon and early evening, and rotationally strip-grazed ones
concentrate their eating activity on the rst meal after a new
pasture strip is allocated, and MINDY simulated it. Despite these
interactions, both cow genotype maintained the same number of
meals. The lack of difference in the daily number of meals matches
previous empirical data (Linnane et al., 2004) where cow genotype
did not affect daily meal number. However, more data are required
to clarify this point and increase condence in model predictions.
Predicted, differences in the number of bites and ingestive
chewing time per bite reect within and between meals uctuations in feeding motivation (Fig. 5). They are consistent with
observed reductions in ingestive chewing with increments in
herbage intake rate for cattle and deer (Greenwood and Demment,
1988; Spalinger and Hobbs, 1992; Gregorini et al., 2009a). Moreover, Orr et al. (1997) and Gibb et al. (1998) and Taweel et al. (2004)
reported fewer chews per bite for sheep and dairy cows as the day
progresses, a pattern also predicted by MINDY. However, MINDY
predicted (Fig. 5) greater bite rates and lower ingestive chewing
time than the daily averages of 53 and 59 bites per min and 0.44
and 0.33 s per bite, reported by Gibb et al. (1998) and Taweel et al.
(2004), respectively, for set stocked dairy cows. Under rotational
grazing, however, bite rate (daily and meal means) fell within the
range of 4065 bites per minute reported in the literature (Rossi
et al., 2004; Gregorini et al., 2009a). Gregorini et al. (2009a) reported
a hunger related decrease in bite rate during meals, from 55 to
39 bites/min. Although the model represented this reduction, predicted bite rates were greater at the beginning of a meal, suggesting
that time per bite may be underestimated (see Eqs. (45 and 27),
respectively). MINDY predicted prehension time of 0.7 s, which
is similar to reported values (Illius and Gordon, 1987). Ingestive
chewing time, the other action determining time per bite, is driven
by bite mass, ber content and feeding motivation. Spatial (grazing

strata) and diurnal changes in toughness of herbage (ber content)
are used in the model to calculate ingestive chewing time, but this
phenomenon requires further development in the model (see Eq.
(30)). Moreover, sward canopy is assumed to be relatively homogeneous, which may add to the potential underestimation of time
per bite.
Predicted daily harvesting and searching times differed between
cow genotypes and grazing management (Table 3). Under rotational grazing both genoypes reduced harvesting time and
increased searching time as the time in the paddock progressed or
herbage was depleted, which is consistent with previous observations for sheep (Ruyle and Dwyer, 1985), blesboks and springboks
(Noville, 1978), wapiti (Hudson and Nietfeld, 1985), beef heifers
(Wade et al., 2006) and dairy cows (Gregorini et al., 2011). These
studies also showed that when these ruminants entered a paddock,
they tended to show higher bite rates than later when the same
pastures had become depleted which is consistent with model
predictions (Fig. 5) for rotationally grazed dairy cows irrespective of genotype. On the other hand, MINDY predicted that under
set stocking grazing management, grazing and harvesting times
(data not shown, but reected by harvesting step rate and herbage
intake rate; Fig. 5) increased and searching step rate decreased
throughout the day. This phenomenon reects the effect of diurnal uctuation of feeding motivation to increase herbage intake
rate as darkness approaches (Gregorini, 2012). MINDY also predicts changes in the dynamics of steps taken while harvesting and
searching within a meal for both grazing methods and cow genotype (Fig. 5) consistent with previous observations of rotationally
grazed dairy cows (Gregorini et al., 2009a) and beef heifers (Wade
et al., 2006; Gregorini et al., 2007a,b) with different hunger levels.
As the meal progresses, the model predicted a decrease in harvesting and an increase in searching step rates. Finally, daily walking
distances while grazing fall within observed ranges (Thomson and
Barnes, 1993; Clark et al., 2010) with surprisingly no substantial difference between cow genotype, but there was a difference between
grazing management (Table 3). Rotational grazing resulted in cows
walking longer distances while harvesting and also when searching.
Despite the novelty of modeling locomotion while grazing, more
data is required in this area.
4.2.2. Ruminating
In MINDY, the length of the rumination bout was modeled as
a function of the size of the ruminal large particle pool which
responds to meal size, rumination rate, and dietary ber content (See Section 2.3). This represents a step forward from Molly
(Baldwin, 1995) where rumination is set to either occur continuously under continuous feeding or periodically using a cosine
function. In the latter case, the proportion of the day during which
rumination occurred was modulated by the amount of dietary ber
so that low dietary ber diets reduced proportional time spent
ruminating. However, the timing of the rumination bouts could not
deviate from the cosine function.
Predicted daily rumination times (Table 3) fall within the range
(387530 min) previously reported by Prez-Prieto and Delagarde
(2012). Daily rumination time was not affected by genotype under
set stocking, which is consistent with the lack of difference among
cow genotypes reported by McCarthy et al. (2007) and Gregorini
(unpublished data). However, under rotational grazing rumination
time was lower for the modern cow type. This phenomenon is consistent with previous observations of Gregorini et al. (2012) and
Gibb et al. (1997) who reported that restricted animals reduced
rumination time as a compensatory behavior to increase grazing
time. Diurnal rumination patterns were affected by cow genotype and grazing management (Fig. 5), which is consistent with
previous observations in dairy cows (Gibb et al., 1997; Gregorini,
unpublished data) and beef heifers (Gregorini et al., 2008b). These

results indicate realistic prediction of rumination time and pattern
by MINDY and its potential to represent rumination time as affected
by cow genotype.
4.3. Rumen function
Ruminal ll and its rate of change within and between meals
were affected by cow type (Fig. 5), indicating that modern, higher
genetic merit (hungrier) cows must either tolerate greater rumen
ll in order to achieve desired intake rates or that rumen ll per
se is not such a strong satiety signal. The strength of satiety signals was set by exponents regulating the effect of each controlling
factor on IHor synthesis (see Eq. (7)). The parameterization of such
exponents (Table 1) indicates that rumen ll is a weaker satiety
signal than rumen VFA (1.02 vs. 10.35, respectively). This greater
strength rumen VFA is concomitant with previous arguments of
Roche et al. (2007) and Sheahan et al. (2011). The effect of rumen
ll in modulating intake pattern should not be discarded though.
Since the sensitivity analysis indicated a signicant effect of rumen
ll on daily grazing and ruination time, as well as number of meals
per day (Table A.3). The differential diurnal pattern of rumen ll,
according to cow genotype and grazing method, can be observed in
Fig. 5. Under set stocking, predicted rumen ll increased throughout
the day to peak at the end of the day, responding to the eating pattern, as previously reported for dairy cows by Taweel et al. (2004)
and Thiago (1988). Under rotational grazing, rumen ll also follows
the eating pattern as reported by Thomson et al. (1985) with sheep.
These results, therefore, evidences realistic prediction of rumen ll
and its effect on intake pattern by MINDY.
The greater satiation power of VFA compared to rumen ll
could also be supported by empirical data showing some compensatory increase of intake when dietary digestibility decrease
(Conrad, 1966). Moreover, increments in milk production are generally explained by an increased rate of removal of VFA from the
rumen and blood, which may explain the increase in feed intake,
as satiety signals from VFA would be reduced, resulting in a longer
meal (Storm et al., 2012). Predicted increments in intake for the
modern cow genotype (Table 3) are consistent with these premises.
Predicted pattern and magnitude of the anabolic hormone changes
with cow genotype and grazing management (Fig. 5). This hormone is a proxy for insulin, which reects the balance of nutrient
absorption and nutrient utilization. Thus, the predicted changes
provide a further link between nutrient demand and intake, supporting intake changes associated with genetic potential for milk
production.
Predicted concentrations of ruminal VFA uctuated during the
day (Fig. 5), which can be related to intake rate and diurnal
changes in chemical composition of the herbage (Taweel et al.,
2004; Gregorini, 2012). From dawn to dusk, herbage accumulates
non-structural carbohydrates (sugars and starch), which dilutes
ber and protein content. This dilution is represented in MINDY.
These diurnal uctuations cause variation in ruminal VFA concentrations from a given meal occurring at different times of the day,
and thus can modulate diurnal intake patterns (Taweel et al., 2004;
Gregorini et al., 2008a,b).
Predicted levels and rate of production of ammonia changed
through the day (Fig. 5), which seemed to be related to grazing
management and therefore meal characteristics (length and intensity) and pattern. Under rotational grazing, predicted increments
in ammonia in the rumen were the greatest for the rst meal of the
day, and were greater in the modern than the older genotype of
cow. Concentration of ammonia in the rumen has been considered
to control herbage intake at the meal level (Conrad et al., 1977;
Miner, 1992; Chapman et al., 2007). The sensitive analysis showed
that meal frequency, length and level of intake are affected by
23
rumen ammonia (Table A.3). However, the parameterized sensitivity exponents in Eq. (7) (Table 1) reect the relative importance of
rumen ammonia in controlling IHor synthesis compared to rumen
VFA. The exponent for VFA is approximately 10 that for ammonia.
Rumen VFA concentration, in particular, has been postulated as a
key factor controlling intake (Illius and Jessop, 1996), which is supported by the reported marked reductions in plasma ghrelin related
to increments rumen fermentation rates as a results of concentrate
supplementation (Roche et al., 2007). Due to rumen fermentation
pattern and rate of production of VFA are related to ammonia availability in the rumen, level of ammonia in the rumen should not be
disregarded and cannot be discarded from the Eq. (7).
Collectively, rumen fermentation end-products and the resulting hormonal changes are the overwhelming modulators of meal
cessation (Pittroff and Soca, 2006; Roche et al., 2008b; Gregorini,
2011). The structure of MINDY allows consideration of this phenomenon and shows it interacting with patterns of intake and
ingestive behavior. Previous modeling efforts, as stated by Allen
(1996); (Allen, 2000), have not been comprehensive enough to
include these more complicated concepts. Therefore, MINDY represents a step forward.
5. Conclusions
The model presented herein, MINDY, makes explicit the functional relationships among direct and indirect controls of feeding
motivation of grazing ruminants. MINDY reproduces the observed
patterns of meals achieving the correct temporal occurrence and
the relative meal lengths of a grazing dairy cow as compared to
those reported in the literature. The models sensitive response to
those functional relationships also allows it to simulate realistic
daily herbage intake and within meal behavior for contrasting grazing environments and cows of different genetic merit. Therefore,
the concepts encoded in the model capture much of the underlying biological mechanisms that drive the diurnal grazing pattern
and ultimately daily herbage intake. This is a considerable advance
in the understanding and modeling of herbage intake and grazing
behavior patterns of free range ruminants.
Estimates of herbage intake and parallel measurements of
ingestive behavior and rumen function of grazing ruminants pose
considerable experimental and technical difculties. As a consequence, advances in knowledge of herbage intake under grazing
conditions have been slow and costly. Therefore, upon completion
of additional testing and evaluation, MINDY can be used to design
and organize experimental programs. The model could also enable
investigators interested in different aspects of the control of intake
and grazing behavior of ruminants (nutrition, physiology, ecology
etc.) to have a common and heuristic tool for mechanistic research.
Thus, MINDY can help to accelerate advances in the knowledge of
the grazing process at low cost.
Acknowledgments
This work was funded by New Zealand dairy farmers thru
DairyNZ Inc. The authors thank Drs. David Chapman and Jeremy
Bryant from DairyNZ (New Zealand) for reviewing this manuscript
and Dave Clark (DairyNZ, New Zealand), Prof. John McNamara
(Washington State University, USA) and Dr. Juan Villaba (Utah State
University, USA) for their useful comments during the development
of this work and writing of the manuscript.
Appendix A.
A.1. Model variables and parameters
Table A.1.
24
Table A.1
Model variables denitions and units.
Symbol
Denition
Value/Unit
ACHT
ActualIR
AHM
AHor
Am
AmCor
BAi
BCS
BCSTarget
BDi
BMi
BR
Chewingfactor
CowHeight
CRi
CurrentStratum
CurrentStratum1
CVFA
DA
Dailydistancewalked
DaylengthP1
Daylight
DayTwlength
DayTwlengthP2
DD
DWH
eLMI
ETHi
ETHini
ETHlim
Fadjustment
FdRat
FSR
GACurrentStratum
GACurrentStratum 1
GAi
GIHor
GrazingSw
HDMI
HGR
Hi
HighChewingMot
HM
HMtotalavail
HMunavail
HSL
iHMtotal
iHMtotalavail
IHor
IHorCor
IHorDeg
IHorRange
IHorSyn
iIHor
kaHor
kAm
kChewfactor
kFdRat
kIHor
kMamCells
kLPSP
kmPref
kRumDM
kVFA
LagDMI
LateFeeding
LM
LMIi
LowChewingMot
LP
MamCellPart
MBDi
MBDsward
MeanLM
MeanSwHeight
Actual chewing time

Actual herbage intake rate of the grazing stratum i
Available herbage mass modulator
Anabolic hormone
Ruminal ammonia concentration
Ruminal ammonia correction factor
Bite area of the grazing stratum i
Body condition score
Body condition score target
Bite depth of the grazing stratum i
Bite mass of the grazing stratum i
Bite rata
Motivation to chew
Animals height to the shoulder
Consumption rate area of grazing stratum i
Upper stratum from the pair strata currently being grazed
Lower stratum from the pair strata currently being grazed
Ruminal concentration of volatile fatty acids
Dental arcade
Daily distance walked
Daylength excluding twilight hours for lactation module
Value representing light intensity
Length of the day including twilight hours
Daylength including twilight hours
Defoliation depth
Distance walked while harvesting
Shape factor
Extended tiller height of the grazing stratum i
Initial extended tiller height
Physical barrier under what cows are not allowed to or are not capable to graze
Adjustment factor to the herbage chemical composition
Herbage intake rate
Number of feeding stations per unit of time
Area harvested at the upper grazing stratum from the pair of grazing strata being grazed at the time
Area harvested at the lower grazing stratum from the pair of grazing strata being grazed at the time
Rates of changes in GSAi due to herbage consumption in grazing stratum i
Motivation to graze
Switch to turn on and off grazing
Daily herbage dry matter intake
Herbage growth rate
Median point height of each grazing stratum i
Constant
Herbage mass
Sum of the herbage mass remaining in each grazing stratum
Unavailable herbage mass
Length of a step while harvesting
Pre-grazing herbage mass
Pre-grazing available herbage mass
Hunger hormone
Scalar
Intake hormone degradation
Constant, Range of IHor
Intake hormone synthesis
Initial IHor
Scalar
Scalar
Scalar
Function adjusting for adiposity and genetic potential
Constant
Scalar (it scales FdRat with genetic potential)
Rate of particle breakdown while ruminating
Scalar to correct PIR(i 1)/PIR(i) to achieve a proper curve shape
Scalar
Scalar
Intake lag function
Adjusting variable to reduce MinIHor
Linear masses of lamina
Linear mass index of each grazing stratum i
Constant
Large particle size pool in the rumen
Number of milk secretor cells in the udder
Mean bulk density of the grazing stratum i
Mean bulk density of the sward
Mean linear mass of the tiller
Half of the ETHini
Days
kg/min
Unitless
Unitless
mmol/L
Unitless
m2
Points
Points
m
kg
Bites/day
Unitless
m
m2 /day
mmol/L
m
m
Unitless
Days
Days
Proportion
m
Unitless
m
m
m
Unitless
kg/day
FS/day
m2
cm2
m2 /day
Unitless
Unitless
kg/day
m/day
m
1, unitless, 31
kg/m2
kg
kg
m
kg
kg
Unitless
1.0, unitless
Unitless
0.02, unitless
Unitless
1.0, unitless
1.0, unitless
0.75, unitless
Unitless
Unitless
6.037, unitless
0.1, unitless
Unitless
Unitless
9.5, unitless
0.11, unitless
Unitless
Days
kg/m
Unitless
0.15, unitless
kg
Unitless
kg/m3
kg/m3
kg/m
m
25
Table A.1 (Continued)

Symbol
Denition
Value/Unit

minimunGSA
MinLPRumntn
MSH
NightMealInter
NightMealTime
Nstrata
Nutrientadjustment
PCHT
PIRCurrent stratum
PIRCurrent stratum+1
PIRi
PREFCurrentStratum
PREFinter
pSTI
PT
Rest
RumDM
Rumntn
SDI
SGR
SM
SSpeedS
SSR
StartIHor
STI
StopIHor
T
TA
TBi
VmIHorSyn
xaHor
xAm
xFdRatLag
xRumDM
xVFA
Area threshold at which hrazing strating has 0 preference

Minimum LP size required to initiate a ruination bout
Momentary speed of harvesting
Length of the last meal of the day
Interval of the last meal of the day and the next meal during the night
Number of sward canopy accessible grazing strata
Adjustment factor to the herbage nutrients
Potential chewing time
Potential intake rate in the upper stratum from the pair strata currently being grazed
Potential intake rate in the lower stratum from the pair strata currently being grazed
Potential herbage dry matter intake rate of the grazing stratum i
Partial preference for the upper stratum from the pair strata currently being grazed
Constant to affect the intercept of the curve of partial preference for current currently being grazed
Momentary average proportion of time searching
Prehension time
Resting (idling) time
Ruminal dry matter load
Rumination time
Distance walked while searching
Sward growth rate
Linear masses of sheath
Momentary average speed while searching
Searching step rate
Constant, trigger point for starting a grazing bout
Searching time
Constant, trigger point for ending a meal
Time of day
Total area offered
Time per bite at the grazing stratum i
Maximum velocity of IHor synthesis
Scalar
Scalar
Scalar (it rescales Roseler et al. (1997) lag function)
Scalar
Scalar
m2
kg
m/d
0.1, unitless
0.044, unitless
Unitless
Days
kg/day
kg/day
kg/day
Unitless
Unitless
Unitless
Days
Days
kg DM
Days
m
m2 /d
kg/m
m/day
Searching steps/day
Unitless
Days
Unitless
Days
m2
Days
Unitless
1.0, unitless
1.12, unitless
0.25, unitless
1.0, unitless
10.0, unitless
Table A.2
Regression parameters relating changes in the nutrient composition of grass with respect to time of day (t, fraction of a day).
Variable name
Constant
Organic matter
Crude protein
Acid detergent ber
Neutral detergent ber
Non-structural carbohydrates
Soluble crude protein
Non-protein nitrogen
Rumen undegradable acid detergent ber
Cellulose
Hemicellulose
88.69
17.20
31.33
50.74
12.32
5.00
22.86
31.96
31.33
16.40
2.93
2.33
10.37
14.14
7.54
0.77
6.14
10.76
9.56
4.57
4.73
41.18
24.20
11.05
50.64
13.28
76.41
25.37
18.59
7.54
6.46
59.93
30.66
14.89
84.43
19.30
108.86
34.85
24.60
39.49
Fadjustment = Constant + a t3 + b t2 + c t.
A.2. Conditional statements

A.2.1. Conditional statements used to ensure proper outputs at
more extreme latitudes
DayTwlengthP2 = 1.0
DayTwlengthP2 = 1.0
If DayTwlengthP2 < 1
If DayTwlengthP2 > 1
A.2.2. Conditional statements used to reduce the StopIHor during

late afternoon and early evening
StopIHor =
StopIHor
NightMult5
value was derived by tting the model to the observations (Gibb

et al., 1998; Taweel et al., 2004).
A.2.3. Conditional statements used to start or stop grazing
Start grazing
GrazingSw = 1
T interMeal = T TMealStop
T interStart = T
End

If IHorstartIHor

If Daylight0

If Daylight < 0 and TInterMealNightMealInter
Stop grazing
If LateFeeding 0
where, StopIHor is the minimum IHor (intake hormone level) level

to stop grazing. NightMult5 is a constant set to 2.0095 to elongate
grazing bout occurring during late afternoon and early evening. This
T interMeal = T TMealStart If GrazingSw = 1

GrazingSw = 0
If IHor StopIHor

T interStop = T
End If Daylight < 0 and TMeal > NightMealTime
0.931x + 0.264 R2 = 0.75

0.154x + 0.269 R2 = 0.37
0.004x + 0.261 R2 = 0.33
0.007x + 0.272 R2 = 0.29
0.018x + 0.206 R2 = 0.98**
0.0528x + 0.2404 R2 = 0.83**
0.0105x + 0.177 R2 = 0.96***
1.799x + 0.095 R2 = 0.93**
2.836x + 0.291 R2 = 0.96**
0.006x + 0.268 R2 = 0.80*
0.016x + 0.177 R2 = 0.96**
3E06x + 0.7085R2 = 0.65***
55.484x + 16.781 R2 = 0.75

9.735x + 17.126 R2 = 0.36
=0.286x + 16.64 R2 = 0.32
0.46x + 17.329 R2 = 0.29
1.18x + 13.163 R2 = 0.98**

3.3782x + 15.268 R2 = 0.83**
0.667x + 11.271 R2 = 0.96***
114.63x + 6.071 R2 = 0.93**
178.4x + 18.494 R2 = 0.96**

0.4109x + 17.083R2 = 0.77*
1.068x + 11.27 R2 = 0.96***
0.0014x + 0.17 R2 = 0.91**
NigthMealInter
NightMealTime
NightMult5
KLPSP
kfdratahor
kfdratAm
kfdratRumDM
kfdratVFA
xaHor
xAm
xRumDM
xVFA
***
**
P < 0.05.
P < 0.01.
P < 0.001.
0.001x + 0.217 R2 = 0.9256**

0.018x + 0.358 R2 = 0.93*
0.082x + 13.817 R2 = 0.92**

1.1x + 22.8
R2 = 0.93*
Daily grazing time (%/d)
Herbage intake (kg DM/d)
Model outputs
VmIHorSyn
kIHor
Variable
1.463x + 0.346 R2 = 0.70

0.008x + 0.319 R2 = 0.13
0.018x + 0.235 R2 = 0.92**
3E05x + 0.004 R2 = 0.51
0.024x + 0.251 R2 = 0.97***

0.023x + 0.320 R2 = 0.32
0.0116x + 0.235 R2 = 0.92***
1.728x + 0.167 R2 = 0.88*
0.467x + 0.333 R2 = 0.75

0.095x + 0.329 R2 = 0.12
0.008x + 0.316 R2 = 0.92*
0.218x + 0.662 R2 = 0.99***
0.0009x + 0.306 R2 = 0.83*

0.016x + 0.420 R2 = 0.96**
Daily rumination time (%/d)
4.300x + 0.362 R2 = 0.91**

0.002x + 0.411 R2 = 0.01
0.035x + 0.587 R2 = 0.94***
3E05x + 0.2847 R2 = 0.57
0.0451x + 0.543R2 = 0.99***

0.076x + 0.439 R2 = 0.88*
0.022x + 0.587 R2 = 0.94***
3.528x + 0.739 R2 = 0.91**
1.3985x + 0.4026 R2 = 0.75

0.058x + 0.401 R2 = 0.05
=0.013x + 0.422 R2 = 0.74**
0.225x + 0.065 R2 = 0.99***
=0.002x + 0.475 R2 = 0.97

0.034x + 0.221 R2 = 0.98**
Daily resting time (%/d)
R2 = 0.75
R2 = 0.73
R2 = 0.07
R2 = 0.56*
R2 = 0.81
R2 = 0.51
R2 = 0.65
R2 = 0.72**
1.0617x + 7.77
2.169x + 3.598
0.212x + 6.276
18.64x + 2.681
181.91x + 2.619
R2 = 0.65
13.4x2 25.7x + 16.6 R2 = 0.55**
0.339x + 6.276
R2 = 0.65
0.0001x + 0.0163
R2 = 0.50
Rest = 0 If FdRat > 0.1

Rumntn = 0
Else If LP > MinLPRumntn
Rumntn = 1
Rest = 0
Else
Rumntn = 0
Rest = 1
1.2x + 0.3
R2 = 0.95**
2.75x + 5.08
R2 = 0.64
0.225x + 2.04
R2 = 0.78**
8.888x + 3.18
R2 = 0.5
200x + 5.98
R2 = 0.93**
13.5x2 + 26.6x 8.9R2 = 0.64**
0.36x + 2.04
R2 = 0.78*
N/A
where, FdRat is intake rate (g/min); Rest (idling) is resting

(min); Rumntn is ruminating (min); LP is pool (kg) of large particles in the rumen; and MinLPRumntn (kg/kg empty BW0.75 )
is the minimum LP size required to initiate a ruination bout.
Empty BW is the weight of the cow without the conceptus
weight.
450x + 5.02
R2 = 0.75
60x + 6.42
R2 = 0.66
8.425x + 42.18
R2 = 0.78
0.4x + 4.6
R2 = 0.45
31.105x + 4.165
57.166x + 2.080
=0.302x + 3.458
0.652x + 3.461
Average intake per meal (kg DM)

R2 = 0.64*
R2 = 0.72
R2 = 0.02
R2 = 0.53
0.054x + 2.15
3.465x + 19.92
0.002x + 4.04
0.175x + 3.46
Meals per day
Table A.3
Responses in model outputs associated with two steps of a 10% change in the value of each model parameter. The regression equation relates variable changes to particular response data.
2.759x + 0.039 R2 = 0.83*

0.2x2 0.4x + 0.3 R2 = 0.65**
0.003x + 0.086 R2 = 0.61
2E07x + 0.0635 R2 = 0.50
0.0165x + 0.119 R2 = 0.93**

0.052x + 0.045 R2 = 0.71
0.002x + 0.086 R2 = 0.61
0.3196x + 0.037 R2 = 0.78*
5.497x + 0.053 R2 = 0.75

0.9121x + 0.030 R2 = 0.76
0.217x + 1s.028 R2 = 0.51
0.005x + 0.059 R2 = 0.34
0.0007x + 0.037 R2 = 0.75*

0.043x + 0.262 R2 = 0.73
Average length per meal (d)
26
where, IHor is the level of the intake hormone, StartIHor is the

minimum IHor level to start grazing, Daylight is the intensity, EatSw
is the switch to turn on and off grazing, NightmealInter is length of
the last meal of the day, MinIHor is the minimum IHor level to stop
grazing and NightMealTime is the interval of the last meal of the day
and the next meal during the night.
A.2.4. Conditional statements used to start or stop rumination
bouts
A.2.5. Conditional statements used to represent foraging

decisions and partial preference for grazing strata
CurrentStratum = CurrentStratum

If GSAcurrentStratum < min imumGSA
KmPREF =
TACurrentStratum PIRCurrentStratum

PIRCurrentStratum /FKmPref
End If PREFCurrentStratum =
If CurrentStratum < NStrata
1 + KmPREF/GSACurrentStratum
PREFCurrentStratum1 = 1 PREFCurrentStratum
IfCurrentStratum < NStrata
Else
KmPREF = 0
(1.0PrefInter)
exp PREF

+ PREFInter
PREFCurrentStratum1 = 0 If CurrentStratum > 1
If CurrentStratum < NStrata
Else
PREFCurrentStratum = 1
where, GSACurrentStratum is the accessible area (m) of the upper grazing stratum from the pair of grazing strata being grazed at the
time. GSACurrentStratum1 is the accessible area of the lower grazing
stratum from the pair of grazing strata being grazed at the time.
PREFCurrentStratum is the relative preference for the upper grazing
PREFCurrentStratum1 is the relative preference for the lower grazing
minimumGSA is a constant that needs to be large enough that the
area consumed per integration interval does not exceed this number. PrefInter, FKmPref and expPREF are also constants. PrefInter is
0.3 affects the intercept of the curve of partial preference for current currently being grazed. FKmPref, 0.2, is a scalar to correct the
ratio PIRi1: PIRi to achieve a proper curve shape. expPREF, 3.0, creates a sigmoid shape to the PREFCurrentStratum (upper stratum of the
pair being grazed at the time).
A.2.6. Conditional statements used to represent the modulatory

effect of time of day
TimeOf DayEffect =
[DawnSwitch (T Dawn) SunriseGIHOR + (Sunrise T )

(NightGIHOR)]
(Sunrise Dawn)
[MorningSwitch (T Sunrise) NoonGIHOR + (Noon T ) (SunriseGIHOR)]
+
(Noon Sunrise)
[AfternoonSwitch (T Noon) SunsetGIHOR + (Sunset T ) (NoonGIHOR)]
(Sunset Noon)
[DuskSwitch (T Sunset) NightGIHOR + (DuskT ) (SunsetGIHOR)]
+
(DuskSunset)
+NightSwitch NightGIHOR
Dawn < Tod Sunrise(i.e.Dawn.DawnSwitch = 1)

Sunrise < Tod Noon(i.e.Morning.MorningSwitch = 1)
Noon Tod Sunset(i.e.Afternoon.AfternoonSwitch = 1)
Sunset < Tod Dusk(i.e.Dusk.DuskSwitch = 1)
Dawn < Tod < Dusk(i.e. Night. NightSwitch = 1)
where, T is time of day (days) and DawnSwitch, MorningSwitch,
AfternoonSwitch, DuskSwitch and NightSwitch indicate the time of
day to the model, and Night, Sunrise, Noon and SunsetGIHOR are
constants, being, 5, 19.5, 21, and 27.8 their respective values scaled
(ScalerTimeofDay) to 15. These values were based on herbage
intake rates reported by Gibb et al. (1998), Gregorini et al. (2007a)
and Orr et al. (1997) in dairy cows, beef heifers and sheep, respectively, under set stocking.
A.3. Diurnal uctuation of herbage chemical composition:
constants
Table A.2
A.4. Local sensitivity analysis
Table A.3
References
Adams, G., Forbes, J., 1981. Additivity of effects of ruminal acetate and either portal
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Ecological Modelling 270 (2013) 1129

Contents lists available at ScienceDirect

A model of diurnal grazing patterns and herbage intake of a dairy cow,

DairyNZ, Ltd., Private Bag 3221, Hamilton, New Zealand

long-term digestive constraints and grazing distribution patterns

P. Gregorini et al. / Ecological Modelling 270 (2013) 1129

none of these models dynamically integrate animal internal state

principles and model parameters are described in this paper. For

P. Gregorini et al. / Ecological Modelling 270 (2013) 1129

et al., 2009) of dairy cows under grazing conditions. Recently the

The model of Forsythe et al. (1995) was used to determine day

Daylight = sin [(T 0.25) 2] 0.5

sin 6/180 + sin Latitude /180 sin (DaylengthP1)

and DaylengthP1, the daylength excluding Twilight hours for the

DaylengthP1 = asin 0.39795 cos 0.2163108 + 2 atan

P. Gregorini et al. / Ecological Modelling 270 (2013) 1129

of ruminal ll and plasma ghrelin in grazing dairy cows. Ruminal

2.2.2. Hunger hormone

where iIHor represents the reference concentration of IHor; set to

1 + CVFA /kVFA kFdRat

where, BCSTarget , is the representation of the target adipose tissue

kMamCells + LagDMI xLag 1

where, VmIHorSyn , CVFA , RumDM, and Am represent, respectively, the

P. Gregorini et al. / Ecological Modelling 270 (2013) 1129

Having dened IHor, trigger points for starting (StartIHor) and

Latefeeding = sin [(T 0.125) 2] 3 0.5

2.3. Diurnal pattern of rumination

sward canopy is relatively homogeneous (horizontal plane) and

[Log (ETHlim ) Log (ETHini )]

For the purpose of calculations of vertical distribution of herbage

Herbage biomass (kg DM/m2 , input to the model) varies through

2.4. Grazing environment and behavior

[(ETHini + MeanSwardHeight) LM] + MeanSwardHeight eLMI (LM SM) / (ETHini + MeanSwardHeight)

P. Gregorini et al. / Ecological Modelling 270 (2013) 1129

While herbage mass is an input to the model, mean bulk density

MBDi = MBDsward LMIi

Temporal changes in the nutrient composition of herbage mass

where, FAdjustmentNDF is dened by Eq. (19). The NDF content is used

where, is BMi given by:

Because BA is dependent on DA (Illius and Gordon, 1987) and

PCHTi = (0.25 NDFi 12.5) BMi

where, PT is based on Illius and Gordon (1987) using empty BW

where, ChewFactor is a variable representing the effect of feeding

where LowChewingMot and HighChewingMot are constants with

2.4.3. Actual motivation to graze

P. Gregorini et al. / Ecological Modelling 270 (2013) 1129

1994; Strubbe and Woods, 2004; Gregorini, 2012). Therefore, actual

where, AHM is the available herbage mass modulator factor at a

(GSAi i MBDi i) BDi

HMunavail = iHMtotal iHMtotavail

iHMtotavail = TA MBDsward (ETHini ETHlim )

where, HMtotalavail is the sum of herbage mass (kg) remaining in

CRi = BRi BAi

where PREFCurrentStratum and PREFCurrentStratum 1 are the relative

where, NutrientCurrentStratum is calculated as described in Section

P. Gregorini et al. / Ecological Modelling 270 (2013) 1129

Having determined PREF, ActualIR, and BM, bite rate (BR,

HSL = CowHeight tan 30

The momentary average speed while harvesting (MSH, m/min)

and the distance walked while harvesting (DWH, m)

Searching time (STI, days) and distance (SDI, m) covered while

where, pSTI momentary proportion of time searching (unitless), and

T interMeal = T TMealStart If GrazingSw = 1

Rest = 0 If FdRat > 0.1