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Vet Clin Food Anim 20 (2004) 447–470

Dry matter intake and energy balance


in the transition period
Ric R. Grummer, PhDa, Doug G. Mashek, PhDb,
A. Hayirli, PhDc
a
Department of Dairy Science, University of Wisconsin–Madison, 1675 Observatory Drive,
Madison, WI 53706, USA
b
Department of Nutrition, University of North Carolina, 2224 McGavran-Greenberg,
Chapel Hill, NC 27599, USA
c
School of Veterinary Medicine, Ataturk University, Erzurum 25700, Turkey

There is no other time in a cow’s life that is more tumultuous than the
transition from late gestation to early lactation. The termination of
pregnancy and initiation of lactation are accompanied by endocrine changes
that exceed those occurring at any other point in time. Concurrent with
shifts in endocrine profile are metabolic changes that facilitate diversion of
nutrients away from maternal stores toward milk synthesis and nurturing of
the newborn. During this critical period, feed intake is at the lowest point
of the lactation–gestation cycle. In practice, we measure feed intake in terms
of dry matter intake (DMI) and commonly use that as an indicator of the
cow’s nutritional status. It seems paradoxical that DMI is lowest when the
demand for nutrients is increasing at the greatest rate a cow will ever
experience.
The decline in feed intake as parturition approaches is not unique to the
bovine, and has been documented for many species. What is unique to the
dairy cow is the unparalleled stress that accompanies the transition from late
gestation to early lactation. As a result of decades of breeding for high milk
yield, within days of parturition it is common for dairy cows to produce 10
times more milk than what is required to satisfy nutrient requirements of
offspring. The resolve to increase efficiency of food production by increasing
production per animal is logical, but it places staggering demands on
periparturient cows. Although cows respond to these demands by increasing
DMI, the response is typically delayed, and consequently, they spend

Corresponding author.
E-mail address: rgrummer@wisc.edu (R.R. Grummer).

0749-0720/04/$ - see front matter Ó 2004 Elsevier Inc. All rights reserved.
doi:10.1016/j.cvfa.2004.06.013
448 R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470

significant time in negative nutrient balance. When negative energy balance


is too severe, it can directly result in metabolic disorders such as fatty liver
and ketosis. Other complications such as retained placenta, displaced
abomasums, and greater susceptibility to infection can be related to the
extent of negative nutrient balances that are experienced by cows during the
transition period. Therefore, to maximize health of periparturient cows, it
seems imperative that we either decrease nutrient demands or enhance
nutrient intake. Because the former is unlikely, it becomes important to gain
a better understanding of feed intake patterns of periparturient cows, factors
that influence feed intake patterns, and relationships between feed intake
and cow health.

Dry matter intake and energy balance during the periparturient period
Because of large variability in moisture content of feeds used in diets fed
to dairy cattle, feed intake is usually expressed on a dry matter (DM) basis.
Dry matter intake depression during the final 2 to 3 weeks before parturition
is well documented. Marquardt et al [1] indicated that DMI decreased 25%
and 52% during the final 14 days of gestation for young (first or second
parity) and aged (third parity or greater) cows. The decline in DMI during
the final 3 weeks of gestation was found to fit an exponential function: DMI
(percent of body weight) at time t = a þ p  e(kt) where a is the asymptotic
intercept at time 1, p is the change in intake (kg) from the asymptotic
intercept until parturition, k is the rate constant determining the shape of
the curve, and t is the day relative to parturition expressed as days pregnant
280 [2]. Data from 172 heifers and 527 cows used in 16 experiments at
eight different universities were pooled to determine coefficients for the
exponential model describing DMI. Dry matter intake, expressed as a
percentage of body weight, at time t equals 1.713  0.688  e(0.344t) for
heifers and 1.979  0.756  e(0.154t) for cows (Fig. 1). Because the DMI
curve is essentially flat at 21 days before parturition, the coefficient
a becomes a reasonable estimate of DMI as animals enter the transition
period. Hence, heifers and cows consume approximately 1.7% and 2.0% of
their body weight at that time. Dry matter intake declines slightly less for
heifers than cows, and reaches approximately 1.3% and 1.4% of body
weight on the day before parturition. The decline in DMI is more gradual
for cows than heifers. The average DMI for the final 21 days before
parturition was 1.88% and 1.69% of body weight for cows and heifers,
respectively. Assuming one knows the proportion of cows and heifers in
a prefresh pen, these values can be used to calculate benchmark intakes for
the pen. Lower DMI, expressed as a % of body weight, for heifers than cows
is somewhat surprising because heifers require nutrients to support growth
in addition to those to support maintenance and pregnancy.
Using the prepartum DMI prediction equations of Hayirli et al [2] and
energy requirement estimates of the National Research Council (NRC) [3],
R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470 449

1.8

1.6

1.4
Heifers
1.2 Cows

1
-21 -19 -17 -15 -13 -11 -9 -7 -5 -3 -1
Day Relative to Calving

Fig. 1. Predicted DMI of heifers and cows during the final 3 weeks of gestation [2].

we predicted energy balance during the final 3 weeks of gestation for heifers
and cows. Assumptions were: 625 kg heifer and 700 kg cow, predicted
average daily gain for heifers according to NRC [3], body condition score of
3.5, and a diet consisting of 40% corn silage, 20% immature legume silage,
23% ground corn, 4% corn gluten meal, 4% heat-treated soybeans, 7.5%
straw, and 1.5% vitamins and minerals (1.68 Mcal net energy for lactation
[NEl]/kg at 13.6 kg DMI/d). The diet used in this example may seem high in
energy density, however, keep in mind that the NEl value was calculated
using the new NRC [3] that allows the NEl content of a diet to increase as
intake is decreased. Results are summarized in Table 1 and Fig. 2.
These data illustrate several important points. First, animals can
experience negative energy balance before parturition, and it is more likely
to occur for heifers than cows. The lower energy balance for heifers can be
explained by their lower DMI and the additional burden of an energy
requirement for growth. Although the data in Table 1 indicates that cows
will not experience negative energy balance at day 279 of gestation, that may

Table 1
Dry matter intake, energy requirements, and energy balance of cows and heifers during late
gestation as predicted by National Research Council
Energy requirement, Mcal NEl
Day of DMI, Energy
gestation kg/day Maintenance Growth Gestation balance, Mcal
Cow 260 13.6 10.2 0 3.5 þ9.2
279 9.3 10.0 0 3.7 þ2.4
Heifer 260 10.7 10.1 1.5 3.5 þ2.5
279 7.8 10.1 1.5 3.7 2.6
Data from National Research Council. Nutrient requirements of dairy cattle. 7th rev.
Washington (DC): National Academy Press; 2001.
450 R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470

Heifer Cow
Heifer + mammary Cow + mammary

10
8
6
4
2
0
-2
-4
-6
-8
255 260 265 270 275 280 285
Day of Gestation

Fig. 2. Theoretical energy balance of a heifer or cow during the final 3 weeks of gestation with
or without considering an energy requirement for mammary growth [3,4]. Assumptions were:
625 kg and 700 kg heifer and cow, predicted average daily gain for heifers according to NRC [3],
body condition score 3.5, and a diet consisting of 40% corn silage, 20% immature legume
silage, 23% ground corn, 4% corn gluten meal, 4% heat-treated soybeans, 7.5% straw, and
1.5% vitamins and minerals (1.68 Mcal NEl/kg at 13.6 kg DMI/d).

not be true if cows experience a greater drop in feed intake than predicted by
NRC [3] or if diets are not as energy dense as that assumed in this example.
If dietary energy density is reduced, it is likely that DMI will be reduced as
well (see factors affecting DMI below). The most recent NRC [3] chose not
to include an energy requirement for mammary growth because insufficient
research has been done to define this requirement. However, if a rough
estimate of 3 Mcal NEl/d is used for mammary requirements [4], cows may
enter into a negative energy balance during the final days of gestation (solid
line with no symbols, Fig. 2). In contrast, when mammary growth is
included, heifers are slightly below zero energy balance during the entire
transition period and may begin to fall significantly below energy require-
ments during the final 10 to 14 days of gestation (dashed line with no
symbols, Fig. 2).
Second, if an animal goes from positive to negative energy balance, it will
be mainly due to the decrease in DMI rather than an increase in energy
requirements for fetal growth. Although increasing requirements for fetal
growth is often cited as a reason that prepartum animals my experience
negative energy balance, energy demands for pregnancy, particularly when
expressed as a percentage of total energy requirements, increase very slightly
(1–2%) during the final 3 weeks of gestation (Table 1). For cows, the 0.2-
Mcal NEl increase in energy required for pregnancy is offset by a decrease in
R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470 451

energy requirements for maintenance. The decrease in energy requirements


for maintenance is associated with decreased DMI [3]. The decrease in
maintenance requirements did not occur for heifers because they grew
during this period, which increased maintenance requirements, thereby
offsetting the effects of reduced DMI.
Reasons for the decline in feed intake before parturition have not been
elucidated but potential mechanisms have recently been reviewed [5,6].
Forbes [7] indicated that the growing fetus might induce space constraints
and restrict rumen volume. Although fetal growth [8] and prepartum DMI
intake [2] have been described as exponential functions, the shapes of the
curves are not consistent. Growth of the fetus is more gradual during the
final trimester of gestation, whereas the drop in DMI does not occur in
earnest until the last few days before parturition. Ruminal water-holding
capacity did not change as cows transitioned from the dry period to
lactation, indicating that physical capacity of the rumen is not the cause of
prepartum DMI depression [9]. If reductions in rumen volume do occur, it is
likely that depression in feed intake could be avoided due to increased rate
of passage of feed particles from the rumen [10,11]. Therefore, it is likely
that factors other than rumen fill must be involved in the regulation of
prepartum feed intake.
Changes in blood concentrations of steroid hormones prepartum are
more consistent with changes in feed intake. We speculated that the surge in
blood estrogen might be responsible for the depression in feed intake before
parturition [12]. Green et al [13] reported a significant negative correlation
(r = 0.46) between feed intake and serum estradiol-17b concentrations in
the blood of late gestation ewes. Injection of estradiol-17b reduced feed
intake in lactating cows [12]; however, administration at day 276 of
gestation to achieve blood concentrations similar to those at parturition did
not depress feed intake [14]. Although biologically less potent than
estradiol-17b, estrone of placental origin is the primary estrogen in blood
at parturition, and the levels are approximately 10-fold higher than those of
estradiol-17b [15]. Dewhurst et al [16] indicated other factors besides
estrogen must be involved with regulating prepartum DMI because the
decrease in DMI is initiated before the rise in estrogen before parturition
[17]. It is likely that multiple factors are involved in the decrease in feed
intake prepartum. Further research is needed to elucidate the role of these
factors that include fat mass, blood metabolites, corticosteroids, leptin,
insulin, gut peptides, cytokines, and neuropeptides [6].

Relationships between prepartum dry matter intake and metabolic disorders


The relationship between prepartum DMI and health and productivity of
cows following parturition has not been clearly defined. Although there are
several studies to suggest a link between DMI and health, there are
numerous studies that indicate there is not one. When reviewing research on
452 R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470

this topic, it is important to remember that large animal numbers are


required to detect significant treatment differences for parameters that
exhibit a binomial distribution, for example, disorders that either occurred
or did not occur. For trials in which animals numbers are insufficient, the
likelihood of falsely concluding there is no treatment difference (referred to
as a type II error in statistics) is increased. When reporting data from
research trials, the probability of committing a type I error (falsely
concluding there is a treatment difference) is reported (as a P-value), but
unfortunately, the probability of committing a type II error is seldom
reported.
Zamet et al [18] monitored 89 Holstein cows through the dry period and
first 30 days of lactation, and categorized cows as being normal or
abnormal. Abnormal (n = 45) cows were those that had one or more
partum or early postpartum disorders diagnosed and treated. Disorders
included dystocia (n = 14), retained fetal membranes (n = 33), parturient
paresis (n = 10), displaced abomasum (n = 7), fat cow syndrome (n = 3),
nephritis (n = 1), and salmonella infection (n = 1) [19]. Dry matter intake
during the final 27-day prepartum decreased from 1.8% to 1.2% of body
weight for normal cows and from 1.8% to 0.9% of body weight for
abnormal cows; DMI was significantly lower for abnormal cows from 3-day
prepartum to 1-day postpartum. Differences in feed intake between the two
groups increased after parturition, and normal cows produced more milk.
This data indicated that predisposition to disorders at and immediately
following parturition may be indicated by reduced DMI prepartum.
Bertics et al [20] suggested that the prepartum decline in DMI may lead to
lipid mobilization from adipose tissue and lead to lipid-related metabolic
disorders such as fatty liver and ketosis. Lipids, released as nonesterified
fatty acids (NEFA) from adipose tissue, are a substrate for oxidation and
are used by many tissues as an energy source during periods of negative
energy balance. Uptake of NEFA by the liver is related to blood flow and
concentration, with estimates being approximately 25% of whole body
uptake [21,22]. Fatty acids taken up by the liver can be partially oxidized to
ketones, completely oxidized to carbon dioxide, or be reesterified to form
triglycerides. The liver has a limited capacity to completely oxidize fatty
acids and the ruminant liver is notoriously slow at exporting triglycerides to
extrahepatic tissues as very low-density lipoproteins [23]. Thus, during
periods of elevated lipid mobilization, oxidation to ketone bodies or storage
as triglyceride (TG) are the predominant fates of NEFA taken up by the
liver. As a result of negative energy balance, cows are predisposed to fatty
liver and ketosis. Cows that were force fed via a rumen fistula during the
prepartum period to maintain DMI had lower liver TG concentration at
1 day after parturition than cows that were allowed to undergo the typical
decrease in DMI before parturition [20]. Plasma NEFA at 2 days before
parturition were 235 lEq/L lower in cows that were force fed, but this
difference was not statistically significant. It should be mentioned that force
R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470 453

feeding cows did not eliminate the rise of NEFA at parturition; therefore,
endocrine and other uncharacterized changes associated with parturition
and lactogenesis are intimately involved with the etiology of fatty liver.
Plasma b-hydroxybutyrate concentrations were not affected by treatment.
However, force-fed cows produced more milk with a higher fat test, so
effects of prepartum treatment on postpartum blood ketone concentration
were confounded by energy output in milk.
Increasing energy and protein density of diets fed to prepartum cows
elevated DMI and lowered plasma NEFA and liver TG [24], indicating that
the beneficial effects of force feeding cows could be obtained through
practical management strategies. A large trial involving cows in commercial
dairy herds indicated that elevated NEFA concentrations during the final
week prepartum were associated with greater incidences of ketosis, displaced
abomasums, and retained fetal membranes [25,26]. Due to the nature of this
trial, DMI were not available. However, if elevated NEFA were a reflection
of lower DMI, this trial supports those of others [18–20,24], and implies that
prepartum DMI has a bearing on subsequent health of cows. There is data
to suggest that in Holstein cows, plasma NEFA become elevated the week
before parturition if DMI falls below 11 kg/d [27–29].
Although data cited above indicates that DMI should be maximized to
prevent metabolic disorders in postpartum cattle, data is available that does
not support this concept. One line of evidence is that heifers consume less
dry matter as a percentage of body weight than mature cows, yet are far less
susceptible to fatty liver [24,30–32]. Although mature cows have slightly
higher peak plasma NEFA concentration at parturition, heifers have an
elevated NEFA concentration leading up to the peak [24,31]. When
examining the relationship between plasma NEFA on day 1 or 2 prepartum
and liver TG at 1 day postpartum, it was obvious that heifers can withstand
higher NEFA concentrations without developing fatty liver than mature
cows (Fig. 3).
A second line of evidence is that cows that are feed restricted also may be
less susceptible to fatty liver. Feed restriction can be achieved either by
limiting feed availability or by increasing dietary fiber. Grum et al [33] fed
a high-fiber diet with low-energy density (1.27 Mcal NEl/kg using the 1989
NRC [34] feed library) or diets with energy density increased by adding fat
(still a high-fiber diet) or by substituting grain for forage (lower fiber diet)
for the entire dry period except the final week before parturition, at which
time they were all adapted to the lactation diet. Cows fed the high-fiber diet
with added fat had the lowest DMI, but did not experience elevated liver TG
at parturition as did cows on the other two treatments. Interestingly,
although cows on the high fiber diet had slightly lower peak plasma NEFA
concentration at parturition (500 versus 675 lEq/L), they had sub-
stantially higher NEFA (400 versus 150 lEq/L) during the weeks lead-
ing up to parturition. This was a scenario quite similar to that described
above for heifers versus mature cows. Because of the experimental design,
454 R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470

Heifers
Cows
Predicted for heifers
5 Predicted for cows
Liver triglyceride (%, wet weight)

0
0 400 800 1200
Prepartum plasma NEFA (µM)

Fig. 3. Relationship between plasma NEFA on day 1 and 2 prepartum and liver triglyceride the
day following calving in heifers and cows (Adapted from VandeHaar MJ, Yousif G, Sharma
BK, Herdt TH, Emery RS, Allen MS, et al. Effect of energy and protein density of prepartum
diets on fat and protein metabolism of dairy cattle in the periparturient period. J Dairy Sci
199;82:1282–95; with permission.)

Grum et al [33] could not delineate if the positive effects of the high-fiber
plus fat diet on liver TG was related to low feed intake or addition of fat to
the diet. In a follow-up study that was designed to examine effects of fat
versus restricted feed intake on liver metabolism of periparturient cows, they
concluded that the beneficial effects on liver TG were due to restricted feed
intake [35].
The results from the study of Grum et al [33] and Douglas et al [35] were
very difficult to reconcile with the force-feeding trial of Bertics et al [20].
Studies on hepatic metabolism by Grum et al [33] indicated that tissue from
cows with the lowest feed intake and highest plasma NEFA concentration
leading up to parturition had the lowest rates of fatty acid esterification and
highest rates of fatty acid oxidation in liver at parturition. This data,
combined with the differences in plasma NEFA and liver TG between
heifers and mature cows [24,31], suggests that livers exposed to elevated
NEFA before parturition may be able to acclimate and adjust their
metabolism to partition NEFA toward oxidation and away from TG
synthesis.
We tested the hypothesis that hepatic tissue acclimates to elevated NEFA
by feeding far-off dry cows either 80% or 160% of energy requirements for 3
weeks and then restricting them to 30% of requirements to induce fatty liver
R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470 455

[36]. The results of this study did not support the hypothesis because prior
energy status did not influence the degree of TG accumulation in the liver
during feed restriction. Perhaps the use of feed restriction in far-off dry cows
was not a suitable model for transition cows. In this trial, we also examined
the effect of parity on susceptibility of fatty liver during feed restriction and
found that, indeed, heifers were less susceptible than mature cows.
Another intriguing similarity between heifers and feed restricted cows is
that they have more consistent DMI during the final 3 weeks before
parturition compared with mature cows that are fed ad libitum diets with
moderate energy concentrations. There is considerable evidence to indicate
that the extent of DMI depression immediately before parturition is
positively related to the level of feed intake [16,32,33,37–41]; an example is
shown in Fig. 4. At 3 weeks before parturition, cows either remained on
a high-fiber diet (49% neutral detergent fiber [NDF], 25% chopped straw)
or were switched to a lower fiber diet (30% NDF, 12.5% chopped straw).
Although cows fed the low fiber diet demonstrated a significant advantage
in DMI during the final 19 day prepartum (2.7 kg/d), they did not have
lower liver TG at 1 day postpartum. Similar results were obtained by Rabelo
et al [32] when feeding diets containing 40% versus 32% NDF before
parturition. This was surprising, considering the positive effects of force
feeding and high DMI previously discussed [20].
The accumulation of data contrary to the conclusions from the force-
feeding trial of Bertics et al [20] has led us [42] and others [43] to reevaluate
the initial interpretation of that study. Perhaps the benefit of force feeding
was not to maximize feed intake, but rather to prevent the decline in feed
intake. This could explain why heifers or cows with relatively low yet
consistent DMI do not experience TG accumulation in the liver at
parturition. Low DMI does not necessarily mean that cows are in negative

16

14
DMI (kg/d)

12

10

6
-20 -15 -10 -5 0
Day Relative to Calving

Fig. 4. Effect of NDF on DMI of late-gestation dairy cows. At 3 weeks before calving, cows
either remained on a high fiber diet (, 49% NDF, 25% chopped straw) or were switched to
a lower fiber diet (, 30% NDF, 12.5% chopped straw) (Data from Minor DJ, Trower SL,
Strang BD, Shaver RD, Grummer RR. Effects of nonfiber carbohydrate and niacin on
periparturient metabolic stats of lactation dairy cows. J Dairy Sci 1998;81:189–200.).
456 R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470

energy balance and mobilizing NEFA. Likewise, a cow with high DMI that
experiences a drop in feed intake may mobilize NEFA from adipose tissue,
even if the drop in feed intake does not put the cow into negative energy
balance. In other words, the most important signal for initiating adipose
tissue lipolysis may be change in intake rather than reaching a threshold
level of energy balance, for examle, zero.
We pooled data from three of our studies with transition dairy cows
[32,40,44] and used regression analysis to test the hypothesis that magnitude
of intake depression is more important than the level of DMI in affecting
metabolic status of cows. Forty heifers and 122 cows were categorized as
having a body condition score greater than or equal to 4.0 (n = 26) or less
than 4.0 (n = 136). The dependent variables in the analysis were plasma
NEFA or liver TG concentration at 1 day postpartum or milk yield or DMI
average for the first 28 days postpartum. Fixed effects in the model were
average daily prepartum DMI (% of body weight) from 21 to 14 days before
expected calving (hereafter referred to as PDMI) or change in daily DMI
between 21 days before expected calving and 1 day before expected calving
(hereafter referred to as CDMI), body condition score, and parity.
Treatment within study was included as a random effect. As an indicator
of magnitude of prepartum DMI, we elected to use average daily DMI from
21 to 14 days before calving rather than from 21 to 1 days before expected
calving. The latter value would encompass change in intake, and we wanted
the two measures to be as independent of each other as possible.
The relationship between PDMI or CDMI and liver TG at 1 day
postcalving is shown in Fig. 5. There was a slight decline in liver TG as
PDMI increased, but the effect was not significant. However, there was
a significant decline in liver TG as the magnitude of intake depression
(CDMI) was decreased. The slope of the lines in each case was not affected
by body condition score or parity; however, the analysis did demonstrate
that heifers have less liver TG than multiparous cows, and cows with a body
condition score equal to or greater than 4 have more liver TG than cows
with less body condition score. As might be expected, similar relationships
were observed for NEFA at 1 day postpartum, that is, CDMI had a greater
and more significant effect on NEFA than did PDMI (Fig. 6). Heifers had
lower plasma NEFA, and overconditioned cows had greater plasma NEFA
than their counterparts at 1 day postpartum.
Interestingly, for DMI and milk yield through 28 days postpartum,
PDMI had a stronger and more significant relationship than did CDMI
(Figs. 7 and 8). If liver TG or plasma NEFA at 1 day after parturition
affects postpartum DMI and milk yield, then one might have expected
similar results for postpartum DMI and milk yield as were obtained for liver
TG and plasma NEFA. This means either elevated liver TG and plasma
NEFA near the time of parturition do not affect postpartum milk yield and
DMI, or the variation in liver TG (9.1  7.2% SD, DM basis) and plasma
NEFA (658  365 lEq/L SD) in these studies were not of sufficient
R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470 457

14 BCS >4 BCS >4


20
Reg. Line Reg. Line
12 Heifers 15 Heifers
10
8 10
6 5
4
0
2
0 -5
0.75 1.25 1.75 2.25 2.75 3.25 -1.7 -1.2 -0.7 -0.2 0.3 0.8 1.3
DMI (% BW) DMI Change (% BW)

Fig. 5. The relationship between prefresh DMI during days 21 to 14 before calving or DMI
change during the final 21 days before calving on TG at 1 day after calving. The solid line
represents the regression generated using data from all animals (P = .32 for DMI, P = .0001
for DMI change). The long dashed line represents cows with a body condition score (BCS) [4
(BCS effect: P = .11 for DMI, P = .07 for DMI change) and the short dashed line represents
heifers (parity effect: P = .0001 for DMI, P = .0001 for DMI change). There were no
interactions with BCS or parity; thus, the slopes of all three lines within a graph are similar
(slope = 1.53 for DMI, 4.96 for DMI change).

magnitude to influence these parameters. The data in Fig. 7 probably


indicates a cow that is genetically predisposed to voluntarily consume
a relatively large quantity of DM does so regardless of what point she is at in
her gestation–lactation cycle.
This preliminary retrospective examination of past trials does suggest
that change in DMI may be an important factor affecting metabolic status
of periparturient dairy cows. Further studies need to be specifically designed
to evaluate the importance of magnitude of DMI versus change in DMI
during the prefresh transition period on health and performance of dairy
cows. Until then, it seems unwise to eliminate the recommendation to
maximize feed intake during the prefresh transition period. However,
perhaps a caveat is in order. If one is to feed for maximum DMI, for
example, by feeding a diet relatively low in NDF, then it would seem
important to avoid situations that may enhance the inherent decline in feed
intake that occurs during this period. Although there is certainly a large
degree of variation among cows in the extent of DMI depression before
parturition, there is evidence to suggest that DMI at 1.3% of body weight is
a fairly typical end point the day before parturition for cows fed a wide
variety of diets [2,45]. Clearly, the higher DMI is above 1.3% of body
weight, the greater the potential to experience a large drop in feed intake.
Consequently, if cows are consuming a high level of DMI (2.0% of body
weight or greater) 2 to 3 weeks before parturition, but are experiencing
severe drops in feed intake before parturition, changing to a diet with higher
NDF to dampen feed intake and flatten the intake pattern before
parturition may be worth considering.
458 R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470

900 900 BCS >4


800 800 Reg. Line
700 700 Heifers
600 600
500 500
400 400
300 300
BCS >4
200 Reg. Line 200
100 Heifers 100
0 0
0.75 1.25 1.75 2.25 2.75 3.25 -1.7 -1.2 -0.7 -0.2 0.3 0.8 1.3
DMI (% BW) DMI Change (% BW)

Fig. 6. Effects of prefresh DMI during days 21 to 14 before calving or DMI change during
the final 21 days before calving on plasma NEFA concentrations at 1 day after calving. The
solid line represents the regression generated using data from all animals (P = .32 for DMI,
P = .0001 for DMI change). The long dashed line represents cows with a body condition score
(BCS) [4 (BCS effect: P = .09 for DMI, P = .09 for DMI change) and the short dashed line
represents heifers (parity effect: P = .006 for DMI, P = .01 for DMI change). There were no
interactions with BCS or parity; thus, the slopes of all three lines within a graph are similar
(slope = 79 for DMI, 172 for DMI change).

Factors affecting prepartum dry matter intake


Factors that affect prepartum DMI can be divided into three main cate-
gories: animal, diet, and management. Although some knowledge has been
gained regarding animal and diet, very little is known about how manage-
ment affects DMI. Much of what we do know about factors affecting feed
intake during the last 3 weeks of gestation comes from the pooled data set
previously mentioned that included 527 cows and 172 heifers that were fed
49 different diets in 16 experiments conducted at eight universities [2,45].
Animal factors
As previously indicated, parity affects feed intake before parturition
(Fig. 1). Heifers eat less as a percentage of body weight, exhibit a more
gradual decrease in DMI, and have a slightly lower degree of DMI
depression. Because of lower DMI and greater nutrient requirements to
accommodate growth, heifers need to be fed a more nutrient dense diet than
mature cows. Heifers are more likely to experience negative energy balance
than mature cows when they are grouped together and fed the same diet.
Breed also affects DMI. French et al [46] developed a prepartum DMI
equation similar to that described by Hayirli et al [2] for Holsteins using 69
multiparous Jerseys fed one of eight diets at three universities. Dry matter
intake at 21 day prepartum was approximately 2.35% of body weight for
Jerseys compared with 1.98% for Holsteins. Prepartum DMI for Jerseys
ranged from 9 to 12 kg/d during the final 3 weeks of gestation and averaged
11 kg/d (2.2% of body weight).
R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470 459

3 3

2.5 2.5

2 2

Reg. Line
Reg. Line
1.5 Heifers 1.5
Heifers
BCS >4
BCS >4
1 1
0.75 1.25 1.75 2.25 2.75 3.25 -1.7 -1.2 -0.7 -0.2 0.3 0.8 1.3
DMI (% BW) DMI Change (% BW)

Fig. 7. Effects of prefresh DMI during days 21 to 14 before calving or DMI change during
the final 21 days before calving on DMI for the first 28 days after calving. The solid line
represents the regression generated using data from all animals (P = .0005 for DMI, P = .47
for DMI change). The long dashed line represents cows with a body condition score (BCS) [4
(BCS effect: P = .0003 for DMI, P = .0001 for DMI change) and the short dashed line
represents heifers (parity effect: P = .04 for DMI, P = .0001 for DMI change). There were no
interactions with BCS or parity; thus, the slopes of all three lines within a graph are similar
(slope = 0.33 for DMI, 0.05 for DMI change).

Although body condition score is ultimately a reflection of management,


it is an animal characteristic that influences DMI. It has been known for
years that postpartum feed intake is decreased in cows that are over
conditioned at parturition [47–49]. The same is true during the prefresh
period (Fig. 9). Cows with thin (body condition score = 1 to 3), moderate
(3.01 to 4) or obese ([4) body condition averaged 1.84%, 1.83%, and
1.68% of body weight during the final 3 weeks of gestation [45]. A
significant body condition score  time interaction indicated that DMI of
obese animals continuously and gradually decreased during the final 3 weeks
of gestation, whereas DMI of animals in thin or moderate body condition
remained relatively constant from 3 to 1 week before parturition, and then
decreased sharply during the final week before parturition. Total DMI
depression during the final 3 weeks was 28%, 29%, and 40% for thin,
moderate, and obese cows, respectively. Because cows in higher body
condition eat less, they are likely to be in a more negative energy balance
compared with cows having lower body condition score fed the same diet.
These changes in feed intake and energy balance in the prefresh period could
potentially contribute to the increased incidences of health and reproduction
problems experienced by over conditioned cows after parturition [50].

Diet factors
We examined the effects of dietary NDF, ether extract (EE), ruminally
undegradable protein (RUP), and ruminally degradable protein (RDP) on
prefresh DMI using our pooled data set [45]. Dietary NDF, RUP, and
460 R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470

45 45
40 40
35 35
30 30
25 25
20 20
15 Reg. Line 15
BCS >4 Reg. Line
10 10 BCS >4
Heifers
5 5 Heifers
0 0
0.75 1.25 1.75 2.25 2.75 3.25 -1.7 -1.2 -0.7 -0.2 0.3 0.8 1.3
DMI (% BW) DMI Change (% BW)

Fig. 8. Effects of prefresh DMI during days 21 to 14 before calving or DMI change during
the final 21 days before calving on milk yield during the first 28 days after calving. The solid line
represents the regression generated using data from all animals (P = .004 for DMI, P = .45 for
DMI change). The long dashed line represents cows with a BCS [4 (BCS effect: P = .34 for
DMI, P = .09 for DMI change) and the short dashed line represents heifers (parity effect:
P = .0001 for DMI, P = .0001 for DMI change). There were no interactions with BCS or
parity; thus, the slopes of all three lines within a graph are similar (slope = 3.48 for DMI, 0.66
for DMI change).

RDP, were categorized according to percentile distributions in 49 diets as


low (lowest 20%), moderate (21%–79%), and high (highest 20%). For EE,
the separation criterion was increased from 20% to 30% to ensure sufficient
number of animals in the low and high group. The animal numbers in each
group were not sufficient to allow for examination of interactions between
animal and dietary factors. Therefore, a second database was created in
which dietary nutrients were categorized as being below or above the mean
value for all 49 diets. Results for mean intakes during the final 3 weeks of
gestation by nutrient category are shown in Table 2.
As expected based on numerous studies [32,33,37–41,51], NDF had the
biggest influence on DMI (Table 2). For cows consuming diets averaging
30%, 42% and 54% NDF, DMI during the final 3 weeks of gestation was
2.03%, 1.68%, and 1.64% of body weight. Further research is needed to
determine if there is a ‘‘break point’’ between 30% and 42% NDF in which
DMI is affected. Considering most prefresh transition diets contain 40% or
greater NDF, it is likely that cows on most commercial farms are not fed to
maximize DMI.
The reason for the increase in DMI when NDF is decreased is puzzling.
In lactating cows, intake of high fiber diets may be limited due to rumen fill
and the inability to degrade fiber fast enough to pass out of the rumen and
make space for additional feed consumption. In this scenario, the cow is not
consuming sufficient energy to meet requirements, so alleviating rumen fill
by decreasing dietary fiber translates into an increase in feed intake. Some
[40,52] have suggested that alleviation of rumen fill might explain the
increase in feed intake when NDF content of dry cow diets is decreased.
R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470 461

1.8

1.6

1.4 BCS ≤ 3.0


BCS > 3.0 to ≤ 4.0
1.2
BCS > 4.0
1
-21 -19 -17 -15 -13 -11 -9 -7 -5 -3 -1
Day Relative to Calving

Fig. 9. DMI of cows and heifers varing in body condition. (Data from French PD, Meyer HH,
James RE, Drackley JK. Dry matter intake prediction equation for nonlactating Jersey cows in
late gestation and breed dry matter intake differences in late gestation. J Dairy Sci
2002;(Suppl 1):44.)

This seems unlikely, because in dry cows, the increase in intake occurs even
if their energy requirements were met before being switched from high- to
low-NDF diets [32,40]. A transient increase in feed intake might be expected
when dry cows are switched to diets lower in NDF, however, the advantage
seems to persist for 3 weeks or longer [32,33,40,52]. The increase in feed
intake occurs even if cows are in adequate body condition (3.5 on a five-
point scale, [32]) and not in need of replenishing body reserves. Estimated
maximum NDF intakes were 0.92% and 0.95% of body weight for dry
primiparous and multiparous dairy cows [53], and similar values have been
reported for prepartum beef cows [54]. Minor et al [40] indicated that NDF
intakes, expressed as a percentage of body weight, were 0.74 and 0.55 for
dairy cows fed diets containing 49% or 30% NDF before parturition.
Therefore, rumen fill probably did not account for treatment differences in
DMI in that study.
Although the pooled data set [45] indicated there were significant effects
of prepartum RUP and RDP on DMI during the final 3 weeks of gestation,
the magnitude of differences were small compared with those observed for
NDF (Table 2). The slight decrease in DMI as RUP is increased may reflect
differences in palatability of dietary ingredients. Interpretation of most trials
examining effects of dietary RUP in prepartum diets is difficult because
changes in RUP are usually accompanied by changes in RDP. Nevertheless,
the overwhelming majority of trials have shown no significant effect of RUP
on prepartum DMI [55–59]. A summary of trials indicated that level of total
crude protein did not affect DMI intake of prefresh cows [60].
Increasing ether extract in prefresh diets adversely affected DMI. The
affect was primarily quadratic, which in this case meant the decrease was
noted when going from diets in the low to medium category. Surprisingly,
this effect occurred with a small change in EE content of the diet (2.0%
462 R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470

Table 2
Influence of dietary nutrients on average dry matter intake during the final 3 weeks of gestation
Average DMI
(% body weight)
Nutrient Average (% DM) for 3 weeks prepartum
NDFa 29.7 2.03
42.5 1.68
53.6 1.64
RUPb 3.5 1.83
4.5 1.80
5.7 1.72
RDPc 8.5 1.73
9.8 1.84
12.9 1.78
EEd 2.0 1.93
3.2 1.71
5.7 1.72
a
Neutral detergent fiber; significant linear (P \ .0001) and quadratic (P \ .0002) effects.
b
Ruminally undegradable protein; significant linear effect (P \ .02).
c
Ruminally degradable protein; significant quadratic effect (P \ .005).
d
Ether extract; significant linear (P \ .04) and quadratic (P \ .0001) effects.
Data from French PD, Meyer HH, James RE, Drackley JK. Dry matter intake prediction
equation for nonlactating Jersey cows in late gestation and breed dry matter intake differences
in late gestation. J Dairy Sci 2002;(Suppl 1):44.

versus 3.2%) and at relatively low levels in the diet. Further inspection for
interactions revealed a strong parity  level of EE interaction. DMI of
heifers and cows were 1.71 and 1.81 of body weight, respectively, when they
were fed diets categorized as below mean EE and 1.25% and 1.70% of body
weight when they were categorized as above mean EE. Thus, DMI of heifers
is more sensitive to changes in EE compared with cows. An explanation for
the interaction is not obvious. Because feeding fat is more common during
lactation than during heifer growth, cows are more likely than heifers to
have been previously exposed to supplemental fat and as a result may be
more tolerant to elevated EE in the diet. Palmquist and Conrad [61] fed
Holsteins and Jerseys diets containing from 3.18% up to 10.8% EE and
reported greater depression of feed intake in Jerseys than in Holsteins.
Although breed is a confounding factor, perhaps frame size influences
animal response to supplemental fat. Research is needed to verify a parity 
dietary fat interaction on prepartum DMI.

Management factors
Of the three categories of factors that may influence DMI (animal, diet,
and management), the least amount of research data is available on
management. To examine the effects of many management factors, farms
rather than cows would have to be the experimental unit, that is, the unit to
R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470 463

which the treatment is applied. This places a heavy emphasis on conducting


research on commercial farms. For logistical reasons, that is seldom done.
Grant and Albright [62] summarized research relevant to feeding
management and behavior of transition cows, but virtually all of the data
was from lactating cows. It is unknown if management factors that influence
feed intake of lactating cows would also affect feed intake of prefresh
transition cows. Assuming they do, then overcrowding, group changes, diet
changes, bunk space, water quality, and so on, may be critical factors
affecting prepartum DMI. Many of the factors just listed are particularly
relevant to dry cows. Frequent housing, grouping, and diet changes are the
norm for cows as they transition from the far-off dry period to lactation.
Dairy expansions result in considerable investment in facilities for the
lactating herd while often times dry cow facilities are not expanded and
become overcrowded. If management factors such as these impact feed
intake, they may have a greater influence in prefresh transition cows than
other animals because they are inherently prone to feed intake depression as
parturition approaches.
With traditional dry cow management schemes, diet changes typically
occur at dry-off, as cows enter the prefresh transition period, and again
immediately after parturition. Cows are fed a high-fiber low-energy diet
during the far-off dry period to avoid excessive weight gain during the dry
period. The prefresh transition period includes a diet change to acclimate
the rumen microorganisms to the low-fiber high-energy diet that is fed after
parturition. Diet changes are usually accompanied by group changes.
Feeding one diet for the entire gestation–lactation cycle would eliminate diet
changes, minimize group changes, and possibly reduce the potential adverse
effects they may have on DMI. However, this alternative feeding strategy
would not work in a traditional 60-day dry cow management scheme
because cows would likely become overconditioned if the diet was high in
energy.
We examined the possibility of reducing the dry period to facilitate
feeding a high-energy lactation-type diet for the entire dry period [63]. There
is abundant research that indicates cows need 50- to 60-day dry periods to
avoid milk production losses the subsequent lactation [64–69]. However,
most research is based on retrospective analysis of farm records. Use of data
from these studies have been criticized because cows with short dry periods
typically calved early, and were not intended to have a short dry period (eg,
cows with twins, cows that spontaneously aborted, cows with incorrect
breeding dates, and so on). Consequently, these cows were not managed for
a short dry period. In contrast, studies that were planned experiments to
examine the effects of shortened dry periods have indicated that 30-day dry
periods may be plausible without significant milk yield losses the subsequent
lactation [70–72].
In our experiment [63], cows were given a 56-, 28-, or 0-day dry period
treatment. Cows on the 56-day dry period treatment had a 28-day far-off
464 R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470

dry period and a 28-day transition period in which a high-fiber low-energy


and moderate-energy diet were fed, respectively. Cows on the 0- or 28-day
dry period were fed a high-energy lactation diet for the entire trial. However,
for the final 28 days of gestation, sodium bicarbonate was removed from the
diet so that it was not excessively cationic, and therefore, less likely to cause
milk fever. Because the dietary scheme was different for cows receiving the
56- or 28-day treatment, those two treatments are a comparison of dry-cow
management strategies rather than a comparison of dry period length per se.
Reducing the dry period from 56 to 28 days to accommodate feeding one
diet for the entire trial did not result in more consistent DMI (Fig. 10). In
fact, the DMI curves are what might be expected based on previous
discussions, that is, dry cows receiving the lower energy density diet for the
last 28 day (cows on the 56-day dry treatment) had lower DMI for most of
the period but less of a drop near the time of parturition than dry cows
fed the high-energy diet (cows on the 28-day dry treatment). Cows that were
on the 0-day dry period treatment had the highest DMI prepartum and
postpartum. The advantage prepartum was expected because cows
continued to milk, and therefore, had greater energy requirements. The
carry over to the postpartum was not expected, particularly because cows
produced significantly less fat-corrected milk than those given 56 or 28 days
dry (33.2 kg/d versus 38.9 or 40.6 kg/d, respectively, for 70 days
postpartum). Liver TG at 1 and 35 days after parturition were 8.4%,
8.8%, and 5.1%, and 8.1%, 5.5%, and 3.1% (DM basis) for cows given 56-,
28-, and 0-day dry periods. Lower liver TG at parturition for 0-day cows
probably reflects mammary removal of NEFA; lower liver TG at 35 days

25

20

15

10

0
-21 -15 -9 -3 3 9 15 21
Day Relative to Calving

Fig. 10. DMI during the transition period of cows given a 56 (), 28 (), or 0 day (6) dry
period. Cows given the 28- or 0-day dry period were fed a high-energy diet before, during, and
after the time frame shown above. Cows given the 56-day dry period were fed a low-energy diet
for the first 28 days of the dry period, a moderate-energy diet during the final 28 days of the dry
period, and a high-energy diet after calving.
R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470 465

after parturition for cows given 28 or 0 days dry probably reflected


differences in energy balance that were observed postpartum. Although data
from this trial did not entirely support our original hypothesis, that
consistency of diet would enhance DMI and health, it did demonstrate
management of dairy cows does influence DMI and liver TG. Our cows
were housed in a stanchion barn at all times. Perhaps if cows had been in
groups and group changes would have accompanied diet changes, then the
concept of consistency benefiting DMI and health may have been realized.

Relationship between prepartum and postpartum dry matter intake


Two independent summaries of previously conducted experiments
indicated that when cows are fed ad libitum, there is a positive relationship
between pre- and postpartum DMI (Mashek et al, unpublished data
presented above) [73]. The correlation (r = .53) between DMI at 1 day
prepartum and 21 days postpartum was highly significant [73]. A similar
correlation was observed between forage DMI at 3 weeks prepartum and
forage DMI during the first 2 to 3 weeks postpartum [16]. Mashek et al
(unpublished; Fig. 7) indicated that for every 1% of body weight increase in
daily DMI at 3 weeks prepartum, there was a 0.33% of body weight increase
in daily DMI during the first 28 days postpartum. The correlation (r)
between DMI at 1 week prepartum and 1 week postpartum was 0.52 for
Holsteins and 0.54 for Jerseys [28]. Removing cows that received more than
38% NDF in their diets increased the correlation coefficient suggesting that
prepartum DMI, when limited by fill, is a less accurate predictor of
postpartum DMI [27,28]. Indeed, several studies have indicated that
restricting feed intake prepartum may increase DMI postpartum [35,39,51].
It is likely that the positive relationship between voluntary prepartum and
postpartum DMI previously noted simply indicates that cows having high
genetic potential for DMI do so regardless of stage of the lactation–
gestation cycle. There is scant evidence that nutritional manipulation to
increase DMI prepartum results in greater DMI postpartum. For example,
increasing DMI during the transition period had very modest [32,74] or no
effect on postpartum DMI [51,52]. Feeding low-fiber high-energy diets for
greater than 3 weeks prepartum may decrease postpartum feed intake if
cows become over conditioned [47,75].
It was previously mentioned that dietary protein had little effect on
prepartum DMI (Table 2). However, increasing dietary protein content of
prepartum diets has been reported to decrease postpartum DMI in some but
not all studies [57,58,76]. Feeding diets containing 12% crude protein (CP)
and 26% of CP as RUP, or 16% CP and either 26%, 33%, or 40% of CP as
RUP, did not influence prepartum feed intake, but postpartum feed intake
through 56 days in milk was lower for cows receiving the 16% CP diets (21.8
versus 19.8, 18.6, and 18.6 kg/d, respectively; [77]). In a follow-up study,
cows were fed prepartum diets containing either 14.1% or 16.2% CP in
466 R.R. Grummer et al / Vet Clin Food Anim 20 (2004) 447–470

which the increase in CP came solely from RUP. Again, no differences in


prepartum DMI were observed, but average daily postpartum DMI through
120 days in milk was 22.7 and 20.8 kg for the 14.1% and 16.2% CP diets,
respectively. This indicated that the effect of protein on DMI in the first trial
was not due to RDP status and nitrogen availability to sustain microbial
growth. Explanations for a carryover effect of prepartum protein feeding are
not obvious. Cows with fatty liver may be compromised in their ability to
convert ammonia to urea [78,79]. Therefore, feeding excess protein before
parturition may result in impaired ability of the cow to detoxify excess
ammonia. It is doubtful these effects would carry out to 120 days in milk.

Summary
It is easy, based on theoretic considerations, to make the argument that
maximizing DMI is important to minimize postpartum complications and
losses in milk production that may be associated with them. However,
research over the past several years provides ample evidence that successful
‘‘passage’’ through the periparturient period is more complicated than
simply maximizing feed intake. Anecdotal evidence from veterinarians and
nutrition consultants also confirm that feeding low-NDF diets to achieve
high prepartum feed intakes during the prefresh transition period does not
necessarily solve fresh-cow problems. Perhaps more important than
maximizing feed intake is to minimize the likelihood of cows experiencing
large drops in feed intake immediately before parturition. Retrospective
analysis of existing data sets indicates that this hypothesis has merit;
research must be conducted to vigorously test it. Until then, it seems
reasonable to try to achieve high DMI, if it can be sustained through
parturition. If it cannot, perhaps a more conservative approach is to limit
voluntary intake by increasing dietary fiber, because data suggests that cows
fed in such a manner experience less dramatic decreases in feed intake as
parturition approaches.
We examined the importance of parity, body condition score, and various
diet components that may influence DMI during the final 3 weeks before
parturition, but they only explained 18% of the variation in intake among
cows [45]. Clearly, there are many other factors that affect intake that need
to be identified. Aspects of farm management that may influence animal
stress need to be investigated, particularly during the prefresh transition
period when cows are inherently prone to reductions in feed intake.

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