Review: Effects of feeding sugars on productivity of

lactating dairy cows

Masahito Oba

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Department of Agricultural, Food and Nutritional Science, University of Alberta, Edmonton, Alberta, Canada
T6G 2P5. Received 29 July 2010, accepted 20 November 2010.
Oba, M. 2011. Review: Effects of feeding sugars on productivity of lactating dairy cows. Can. J. Anim. Sci. 91: 37
46. Sugars
are water-soluble carbohydrates that are readily available in the rumen. Although sugars ferment faster than starch or fibre
in the rumen, the rates of disaccharide hydrolysis and monosaccharide fermentation vary greatly depending on the type of
sugar and rumen environment. Despite rapid fermentation in the rumen and their potential to provide greater fermentable
energy to enhance microbial protein production, feeding sugars in place of dietary starch sources may not decrease rumen
pH or improve N utilization efficiency and milk protein production in dairy cows. However, feeding high-sugar diets often
increases dry matter intake, butyrate concentration in the rumen, and milk fat yield. These nutritional characteristics of
sugars may allow us to use high-sugar feedstuffs as an alternative energy source for lactating dairy cows to increase dietary
energy density with reduced risk of rumen acidosis, but there is little evidence in the literature to indicate that the
synchrony of rumen fermentation would be enhanced by feeding high-sugar diets with high soluble protein. Greater
butyrate production from feeding high-sugar diets is expected to enhance proliferation of gut tissues, but its physiological
mechanisms and effects of butyrate metabolism on overall productivity of dairy cows warrant further investigations.
Key words: Dairy cows, ruminants, rumen fermentation, sugars, sucrose, lactose
Oba, M. 2011. Les sucres dans lalimentation et leur incidence sur le rendement des vaches laitie`res en lactation. Can. J. Anim.
Sci. 91: 37
46. Les sucres sont des hydrates de carbone hydrosolubles quabsorbe facilement le rumen. Bien quils fermentent
plus rapidement que lamidon ou la cellulose, la rapidite de lhydrolyse des disaccharides et de la fermentation des
monosaccharides varient considerablement selon la nature du sucre et les conditions dans le rumen. Malgre une
fermentation rapide dans le rumen et la liberation eventuelle dune quantite superieure denergie qui favorisera la production de proteines unicellulaires, lusage de sucres au lieu damidon dans les aliments ne reduira pas forcement le pH du
rumen et nameliorera pas necessairement lassimilation du N ni la production de proteines dans le lait des vaches. En
revanche, une ration riche en sucres accro tra souvent lingestion de matie`re se`che, la concentration de butyrate dans le
rumen et le rendement du lait en matie`re grasse. Ces particularites nutritionnelles des sucres pourraient deboucher sur
lusage daliments du betail riches en sucres en tant que source denergie de rechange pour les vaches laitie`res en lactation,
grace a` une hausse de la quantite denergie issue des aliments sans que le risque dacidose du rumen sen trouve accru.
Malheureusement, peu detudes indiquent que le recours a` une ration riche en sucres et en proteines hydrosolubles
ameliorerait le synchronisme de la fermentation du rumen. On sattend a` ce que la plus forte production de butyrate derivant
dune alimentation riche en sucres favorise le developpement des tissus intestinaux, mais il faudrait entreprendre dautres
etudes pour preciser la physiologie et les consequences du metabolisme du butyrate sur la productivite des vaches laitie`res.
Mots cles: Vaches laitie`res, ruminants, fermentation du rumen, sucres, sucrose, lactose

Carbohydrate fractions contribute to approximately

70% of dry matter (DM) in the diets for lactating dairy
cows. They greatly vary in digestion rate and fermentation end products in the rumen, subsequently affecting
nutrient utilization by animals. Fibre ferments slowly in
the rumen and often limits voluntary feed intake by physical fill (Allen 1996). Starch ferments faster than fibre
and can decrease rumen pH, resulting in more propionate production in the rumen (Owens and Goetsch 1988).
Although high-starch diets increase the energy density,
the excess propionate production may cause hypophagia
(Allen 2000) and the low rumen pH often causes milk fat
depression (Kalscheur et al. 1997). Therefore, it is important to understand the unique characteristics of each
carbohydrate fraction to optimize milk production of
dairy cows, but our understandings of sugar nutrition
and its effects on animal metabolism are limited.
Can. J. Anim. Sci. (2011) 91: 37
46 doi:10.4141/CJAS10069

Sugars are water-soluble carbohydrates, and include

disaccharides, such as sucrose, lactose, and maltose, and
monosaccharides, such as glucose, galactose, and fructose. Sugar is generally considered to be a carbohydrate
fraction that ferments very quickly in the rumen. The
Cornell Net Carbohydrate and Protein System assumes
a fermentation rate of 300% h 1 for sugars (Sniffen et al.
1992). Although dairy diets are not usually formulated
for a specific sugar concentration, almost all feedstuffs
used in dairy diets contain some form of sugar, and some
feedstuffs have a very high sugar content. Starch and
Abbreviations: BHBA, b
hydroxybutyrate; DM, dry matter;
DMI, dry matter intake; NDF, neutral detergent fibre; NEFA,
non-esterified fatty acids; OM, organic matter; RDP, rumen
degradable protein; SIP, soluble intake protein; WSC, water
soluble carbohydrates
37

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CANADIAN JOURNAL OF ANIMAL SCIENCE

fibre are primary carbohydrates fed to dairy cows, but

sugars can be good alternative energy sources, and their
impacts on rumen fermentation and animal productivity
are of interest among ruminant nutritionists. Therefore,
the objectives of this paper are to review the impacts
of sugars on rumen fermentation, and to evaluate the
effects of feeding sugars and high-sugar feedstuffs on the
productivity of dairy cows.

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ROLES OF SUGARS IN RUMEN

FERMENTATION
Volatile Fatty Acid Profile
As the type of fermentation end-products affects nutrient utilization by animals, it is important to understand
how sugars alter the volatile fatty acid profile. Many in
vitro studies in the literature have indicated that feeding
sugars can increase butyrate production in the rumen;
for example, it has recently been shown that the substitution of sucrose for corn starch (Vallimont et al. 2004) or
hay (Ribeiro et al. 2005) increased the molar proportion
of butyrate. However, the effects of feeding sugar on
rumen fermentation in vivo have been considerably
variable; the molar proportion of butyrate in the rumen
fluid in vivo was increased (Kellogg and Owen 1969a, b),
not affected (Sannes et al. 2002; Broderick et al. 2008;
Penner and Oba 2009), or tended to be decreased
(McCormick et al. 2001) by partial replacement of grain
with sucrose.
Type of sugars also affects the volatile fatty acid
profile in rumen fluid; the molar proportion of butyrate
in vitro was increased with the addition of glucose,
fructose, or galactose, but not with the addition of xylose or arabinose (Sutton 1968). In sheep fed grass silage, feeding xylose and fructose increased the molar
proportion of propionate in rumen fluid, while feeding sucrose and lactose increased the molar proportion
of butyrate (Chamberlain et al. 1993). Feeding lactose
in place of ground corn increased butyrate, but decreased propionate concentration in the rumen fluid of
steers (Schingoethe et al. 1980) or lactating dairy cows
(DeFrain et al. 2004). Similarly, feeding milk in place
of concentrate mix increased butyrate, but decreased
propionate concentration in rumen fluid (Doreau et al.
1987). Heldt et al. (1999) noted that feeding glucose,
fructose, or sucrose in place of starch increased valerate
concentration in the rumen. However, Broderick et al.
(2008) showed that feeding sucrose in place of starch
did not increase valerate concentration linearly but
quadratically, peaking at 2.5% of sucrose addition on
a DM basis, and others reported that it was not affected
by feeding sucrose in vivo (Kellogg and Owen 1969a;
Kellogg 1969) or in vitro (Vallimont et al. 2004). Contrarily, valerate concentration was consistently increased
by feeding lactose (Bragg et al. 1986; Doreau et al. 1987;
DeFrain et al. 2004).
Inconsistencies observed for effects of sugars on
rumen fermentation can be also attributed to feedstuffs

that were replaced by sugar. Feeding sucrose to wethers

fed lucerne hay cubes increased the concentration of propionate as well as butyrate in the rumen fluid (Sutoh
et al. 1996). However, this observation can be attributed to greater dietary non-fibre carbohydrate intake
rather than specific effects of sugar fermentation. Studies
where sugar partly replaced dietary starch showed that
the molar proportion of propionate was decreased
(Schingoethe et al. 1980; Heldt et al. 1999; DeFrain
et al. 2004) or not affected (Kellogg and Owen 1969a, b;
Vallimont et al. 2004).
Furthermore, although some studies reported that
molar proportion of butyrate in rumen fluid was not
affected by feeding sugars, it should be noted that
butyrate production in the rumen is not same as butyrate
concentration because the concentration is a function
of production, absorption, and passage of butyrate. Because absorption of butyrate is faster than that of acetate
or propionate (Leek 1993), butyrate concentration in
rumen fluid, either as molar-% or mM, likely underestimates the actual butyrate production. Penner et al.
(2009) and Penner and Oba (2009) reported that feeding sucrose in place of cracked corn tended to increase
rumen pH without affecting butyrate concentration in
rumen fluid, but the possibility of greater butyrate production for sucrose treatment cannot be excluded. Thus,
effects of feeding sugar on actual fermentation acid
production should be evaluated to determine if butyrate
production in the rumen is affected by feeding sugars.
Fermentation of sugars in the rumen initiates with the
hydrolysis of disaccharides to monosaccharides, followed by the fermentation of monosaccharides, but
the rates of disaccharide hydrolysis vary greatly among
different types of sugars. Weisbjerg et al. (1998) reported
that the hydrolysis rate of sucrose is faster than that of
lactose (1404 vs. 248% h1) in animals that are not
adapted to diets supplemented with sucrose or lactose,
and that lactose hydrolysis is greater in animals adapted
to a high lactose diet compared with control animals
or those adapted to a high sucrose diet (540 vs. 248
and 204% h1, respectively). These results indicate that
different microbial organisms may be involved in the
hydrolysis of different types of disaccharides or that
microbial adaptation may be required for the hydrolysis
of specific disaccharides. Monosaccharides also vary
in their rate of fermentation. Sutton (1968) evaluated
five types of sugars (glucose, fructose, galactose, xylose,
arabinose), and reported that glucose and fructose were
almost completely fermented during 2 h of in vitro incubation, while approximately 50% were fermented for
galactose (a component of lactose), xylose, and arabinose (both are five-sugar monosaccharides in hemicelluloses). These results are consistent with those of Cullen
et al. (1986), who reported greater lactate production
and lower pH from 12 h of in vitro fermentation of
glucose, fructose, or galactose compared with that of
xylose or arabinose. Both Sutton (1968) and Cullen
(1986) used rumen contents collected from animals that

OBA * FEEDING SUGARS AND DAIRY COWS 39

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were not adapted to a high-sugar diet for their in vitro

studies. Contrarily, Weisbjerg et al. (1998) reported that
the rate of glucose fermentation varies from 422 to
738% h1 depending on whether animals are adapted
to a high-sugar diet, indicating that variations observed
in the fermentation rate of sugar is at least partly
attributed to variations in microbial activities as well as
substrate type. Sucrose is expected to ferment completely in the rumen due to the rapid hydrolysis and
subsequent fermentation of monosaccharides, but the
minor portion (i.e., B5%) of lactose may escape rumen
fermentation and reach the duodenum, and be available
for enzymatic digestion (Weisbjerg et al. 1998).
Rumen pH
Due to the rapid fermentation of sugars relative to the
other carbohydrate fractions, rumen pH is expected to
be lower for diets containing sugars. However, many
studies in the literature show that rumen pH is not
affected when dietary starch sources are partly replaced
by sucrose (Sutoh et al. 1996; McCormick et al. 2001;
Broderick et al. 2008) or lactose (Schingoethe et al. 1980;
DeFrain et al. 2004). Furthermore, some studies report
that rumen pH increases (Chamberlain et al. 1993; Heldt
et al. 1999) or tend to increase (Penner et al. 2009;
Penner and Oba 2009) with the partial replacement
of dietary starch sources with sugar. Kellogg (1969)
showed that feeding sucrose in place of ground milo
decreased rumen pH. This is one of few in vivo studies
that reported lower rumen pH for a diet containing
sugar in place of dietary starch sources, but their finding
might be attributed to the extremely low dietary forage
allocation; Kellog (1969) fed a diet containing only 10%
forage, while experimental diets used in the other studies
mentioned above contained at least 30% forages on a
DM basis. Collectively, there is little evidence in the
literature to support the concept that increasing dietary
sugar concentration decreases rumen pH, suggesting
that the effects of digestion rate on rumen fermentation
cannot be compared among different carbohydrates
(Hall 2004).
It is not known why feeding sugar potentially increases rumen pH despite its rapid fermentation in the
rumen. However, there are several theories to explain
high rumen pH for animals fed diets containing sugar in
place of starch. One possible explanation is that sugar
provides less carbon compared with starch for fermentation acid production per unit of mass (Hall and Herejk
2001). In addition, if greater dietary sugar supply increases the rate of passage (Sutoh et al. 1996) or production of microbial mass (Ribeiro et al. 2005), less
organic matter (OM) would be available for fermentation acid production (Allen 1997). Microbial glycogen
synthesis from sugars is another possible explanation;
microbes can convert sucrose to glycogen as a short-term
energy storage (Hall and Weimer 2007), which temporarily reduces fermentation acid production in the rumen,
possibly contributing to higher rumen pH. Masson and

Oxford (1951) reported that glucose, fructose, and

sucrose were utilized for glycogen synthesis by holotrich
ciliates, but other sugars such as galactose, mannose,
xylose, arabinose, lactose, cellobiose, and maltose were
not extensively utilized for glycogen synthesis (Oxford
1951). Furthermore, greater butyrate production in the
rumen from feeding sugar (Vallimont et al. 2004; Ribeiro
et al. 2005) would decrease proton production per unit
of ruminally degraded OM compared with acetate or
propionate production as 1 mole of hexose ferments to
1 mole of butyrate, or 2 moles of propionate or acetate
(Owens and Goetsch 1988). In addition, as discussed
above, many studies showed that feeding sugars can
increase the concentration of valerate in the rumen,
which further contributes to less proton production per
unit of OM fermented in the rumen.
The type of sugar also affects rumen pH. Weisbjerg
et al. (1998) measured the changes in rumen pH after a
single dose of sugar and reported that a sucrose dose
decreased the rumen pH to the greatest extent followed
by a dose of glucose or lactose. This supports the similar
observations made by Chamberlain et al. (1993), and is
consistent with a slower hydrolysis rate for lactose
compared with sucrose (Weisbjerg et al. 1998) and less
fermentation for galactose compared with glucose or
fructose (Sutton 1968). Heldt et al. (1999) reported that
steers fed glucose at 0.3% of body weight per day (DM
basis) had lower rumen pH than those fed fructose;
however, others reported no differences in fermentation
rate between glucose and fructose (Sutton 1968) or lactate production (Cullen et al. 1986). The impacts of sugar
fermentation on rumen pH warrant further investigation. Rapid disappearance of sugar per se does not necessarily mean extensive fermentation acid production
and low rumen pH, and sugars might be used as an
alternative energy source that reduces risk of rumen
acidosis in ruminant diets.
Microbial Protein Production
Rapid fermentation of OM is expected to increase
microbial protein production in the rumen (Nocek and
Russell 1988). Feeding rapidly fermentable carbohydrates is expected to capture more ruminally degradable
N (Sniffen et al. 1992), and there are some reports that
dietary inclusion of sucrose decreased ammonia-N concentration in rumen fluid (Chamberlain et al. 1993;
Sannes et al. 2002; Broderick et al. 2008), but others
show no effect of sucrose supplementation on rumen
ammonia concentration in vitro (Vallimont et al. 2004)
or in vivo (McCormick et al. 2001; Penner and Oba
2009). The discrepancy may be attributed to the difference in fermentability of basal diets; diet fermentability before sucrose supplementation may not have
limited ammonia-N utilization in some studies. It is
noteworthy that feeding lactose decreased ruminal
ammonia concentration in prepartum cows (DeFrain
et al. 2006), but not in lactating cows fed more fermentable diets (DeFrain et al. 2004; DeFrain et al. 2006).

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Feeding high-sugar diets may decrease ammonia-N

concentration in rumen fluid if readily fermentable
carbohydrates limited microbial protein production,
but marginal effects of sugar addition on the ammonia-N concentration may be negligible if basal diets are
already high in fermentability in the rumen.
The effects of feeding sugar on microbial protein
synthesis are not consistent in the literature. Glucose
infusion increases microbial N in cattle fed grass silage
(Rooke et al. 1987). Similarly, ruminal infusion of
sucrose increased microbial protein production in nonlactating cows fed grass silage (Kim et al. 1999b, 2000)
and wethers (Kim et al. 2005). Although ruminal infusion of sugars enhanced microbial protein production,
it should be noted that sugar infusion increased total
fermentable carbohydrate intake, which confounded the
specific effects of sugar. Partial replacement of barley
grain with dextrose increased bacterial N flow to the
duodenum (Piwonka et al. 1994). However, others reported that bacterial N was decreased (Sannes et al.
2002) or not affected (Broderick et al. 2008) by dietary
inclusion of sucrose. These discrepancies can be partly
explained by the difference in experimental approaches;
what was actually replaced by sugars. Dietary sugar inclusion may not increase microbial protein production if
it dilutes dietary starch. In vitro studies in the literature
support this speculation; microbial protein production
was increased when sucrose replaced alfalfa hay (Ribeiro
et al. 2005), but decreased (Hall and Herejk 2001) or not
affected (Vallimont et al. 2004) when sucrose replaced
starch. Sugar may be less efficient at increasing microbial
protein synthesis compared with starch, partly because
sugar provides less carbon than starch on a mass basis
(Hall and Herejk 2001). In addition, a faster rate of fermentation of sugar relative to starch may increase energyspilling, which decreases energy available for microbial
growth (Strobel and Russell 1986). These findings were
not in agreement with those of Chamberlain et al. (1993),
who reported greater microbial protein production for
sheep supplemented with sucrose compared with those
supplemented with starch.
Nutrient Digestibility
Sucrose supplementation decreased the rate of neutral
detergent fibre (NDF) digestion (Huhtanen and Khalili
1991) and the activities of carboxymethylcellulase and
xylanase extracted from microbes (Huhtanen and Khalili
1992). However, the negative effects of sugars on NDF
digestion may not be caused by low rumen pH, since the
addition of glucose decreased the rate of NDF digestion
in vitro even if the pH of media was maintained above
6.2 (Piwonka and Firkins 1993). Piwonka and Firkins
(1996) showed that the end products of glucose fermentation, proteinaceous inhibitors rather than fermentation
acids, may be responsible for reduced NDF digestion in
vitro. However, feeding sucrose in place of dry cracked
corn at 4.4 to 4.7% of dietary DM did not affect total
tract digestibility of any nutrients in lactating dairy cows

(Penner and Oba 2009; Penner et al. 2009). When sucrose

replaced starch at up to 7.5% of dietary DM, a quadratic
effect was observed for ruminal NDF digestibility, with
a peak at 5.0% sucrose in the diet (Broderick et al. 2008).
Furthermore, Heldt et al. (1999) reported that sugar supplementation (0.3% of BW d1) increased NDF digestibility compared with the control (a high-forage diet) or
starch supplementation. Therefore, the negative effects
of sugar fermentation on NDF digestion may not be
expected as long as the addition of sucrose is limited to a
reasonable level, and more research is warranted to
identify the threshold of sugar addition and interacting
factors that negatively affect NDF digestibility.
IMPACTS OF SUGARS ON PRODUCTIVITY OF
RUMINANTS
Dry Matter Intake
The inclusion of sugar in diets of lactating dairy cows
often increases dry matter intake (DMI). Broderick
(2008) reported that DMI increased linearly as sucrose
supplementation increased to 7.5% of dietary DM.
Similarly, Penner and Oba (2009) reported that feeding
sucrose at 4.7% of dietary DM increased DMI in early
lactating cows. Furthermore, DeFrain et al. (2004)
showed that DMI tended to increase linearly as lactose
inclusion increases up to 14.2% of dietary DM. Dairy
cows likely recognize and prefer the sweet taste over the
other primary tastes (Nombekela et al. 1994). Although
a sweetened diet is preferred if animals are allowed to
choose a diet to consume (Murphy et al. 1997), sucrose
addition did not increase DMI of dairy cows in early
lactation (Nombekela and Murphy 1995), possibly because sucrose inclusion at 1.5% of dietary DM may not
be high enough to elicit the DMI response. Although
some researchers reported that DMI was not affected by
supplementation with sucrose (McCormick et al. 2001;
Penner et al. 2009) or lactose (DeFrain et al. 2006), to the
authors knowledge, there is no report in the literature
showing the negative effects on DMI of feeding sugars.
Nutrient Utilization
Feeding high-sugar diets is often associated with greater
butyrate concentration in the rumen. Butyrate is extensively metabolized by ruminal epithelial cells (Kristensen
2005), and its effects on energy metabolism of ruminal
epithelia and gut proliferation have been extensively
studied. Sakata and Tamate (1978) demonstrated that
butyrate increases cell proliferation, but it seems to be
mediated by systemic insulin (Sakata et al. 1980), IGF-1
(Shen et al. 2004), or the autonomic nervous system
(Sakata and Engelhardt 1983) rather than local effects
of butyrate metabolism within ruminal epithelia (Wang
and Jiang 2010). Nonetheless, it is noteworthy that
greater butyrate concentration in rumen fluid was associated with greater feed efficiency of beef cattle (Guan
et al. 2008). The effects of feeding sugar and subsequent
enhanced butyrate absorption on gut proliferation and

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OBA * FEEDING SUGARS AND DAIRY COWS 41

overall nutrient metabolism in ruminants warrant

further investigation.
The effects of feeding sugars on the efficiency of N
utilization are not consistent in the literature. Sucrose
supplementation often decreased urinary N excretion
(Sutoh et al. 1996; Sannes et al. 2002; Broderick et al.
2008) and increased N retention (Sutoh et al. 1996).
However, others reported that partial replacement of
corn grain with sucrose increased (McCormick et al.
2001) or tended to increase (Penner and Oba 2009)
plasma urea N. In addition, efficiency of N utilization for
milk protein production decreased linearly as sucrose
inclusion increased (Broderick et al. 2008). Furthermore,
milk protein concentration was decreased by adding
sucrose to TMR at 1.5% of dietary DM (Nombekela and
Murphy 1995). These data suggest that a partial replacement of dietary starch with sugars does not necessarily
improve N utilization.
Contrarily, the effects of feeding high-sugar diets on
milk fat production and energy metabolism are more
consistent in the literature. Although, to the authors
knowledge, none of the previous research reported that
sugar supplementation increased milk yield, several
studies reported that cows fed high-sugar diets increased
(Broderick et al. 2008; Penner and Oba 2009) or tended
to increase (Nombekela and Murphy 1995) milk fat
yield. These responses may partly be attributed to
reduced trans-fatty acid production in the rumen. Incomplete biohydrogenation of unsaturated fatty acids is
a primary cause of milk fat depression (Shingfield and
Griinari 2007), but Ribeiro et al. (2005) demonstrated
that biohydrogenation of unsaturated fatty acids decreased linearly with sucrose addition in continuous
culture media. In agreement with their findings, Penner
and Oba (2009) showed that feeding a high-sugar diet
decreased C18:1-trans fatty acid concentration in milk
fat, and tended to increase milk fat yield.
Another consistent effect associated with feeding
sugars is the increase in plasma b
hydroxybutyrate
(BHBA) concentration (DeFrain et al. 2004; Penner
and Oba 2009). The BHBA comes either from incomplete fatty acid oxidation in the liver or butyrate metabolism in the digestive organs. As Penner and Oba (2009)
reported greater plasma NEFA concentration for cows
fed a high-sugar diet in early lactation, the possibility
of greater NEFA mobilization and subsequent greater
ketogenesis in the liver cannot be excluded. However, in
their study, cows fed the high sugar diet increased DMI
and energy intake, and did not have greater negative
energy balance compared with cows fed the control diet.
Thus, it is not clear why the high-sugar diet increased
plasma NEFA concentration. Contrarily, feeding lactose
to cows either in late lactation (DeFrain et al. 2004) or
early lactation (DeFrain et al. 2006) increased plasma
BHBA concentration without affecting plasma NEFA
concentration, and cows fed high-lactose diets decreased
liver lipid content by approximately 40% (DeFrain et al.
2006). Therefore, greater plasma BHBA concentration

for cows fed high-sugar diets is not likely to subject

animals to ketosis, but the source of BHBA in cows fed
high-sugar diets is still a matter of debate.
UTILIZATION OF HIGH-SUGAR FEEDSTUFFS
Forages
Forages contain moderate amount of sugars, and the
sugar content can be greatly affected by forage variety
(Humphreys 1989; Berthiaume et al. 2010). Berthiaume
et al. (2010) reviewed research in the literature that
evaluated ryegrass cultivars containing greater watersoluble carbohydrates (WSC; i.e., sugars), and found
that animals fed high-WSC ryegrass cultivars increased
microbial N flow (Lee et al. 2002), microbial efficiency
(Merry et al. 2006), efficiency of N utilization (Miller
et al. 2001), and average daily gain (Lee et al. 2001)
compared with those fed control ryegrass. Similarly,
Berthiaume et al. (2010) reported that alfalfa cultivars
with greater concentrations of non-structural carbohydrate (particularly for sucrose and starch) increased
microbial N flow in continuous culture.
The sugar content of forage is also affected by maceration and harvest time (AM or PM). Plants accumulate
sugars during the daytime by photosynthesis, and consume sugars by respiration during the night-time. Thus,
forages harvested in late afternoon are expected to have
greater sugar content compared with those harvested in
the morning. Tall fescue hay cut at sundown contained
more monosaccharides and disaccharides compared with
that cut at sunup (Fisher et al. 1999). Similar observations were made for alfalfa hay (Fisher et al. 2002) and
alfalfa baleage (Brito et al. 2008). Intensive mechanical
conditioning of forages, such as maceration, also affect
their sugar content (Savoie 2001); maceration is expected
to maintain high sugar content of forages by decreasing
wilting time and minimizing respiratory loss, but may decrease sugar content of forages to a greater extent under
poor wilting conditions (e.g., rain or high humidity).
High-sugar forages are expected to increase microbial
protein production in the rumen and animal productivity. Trevaskis et al. (2001) reported that grasses cut in
late afternoon contained more WSC than grasses cut in
the early morning, and that sheep fed afternoon-cut forages excreted more allantoin in urine indicating greater
microbial protein production in the rumen. Similarly,
dairy cows fed alfalfa baleage cut at sundown, which
contains greater WSC, increased milk yield, efficiency
of N utilization (milk N/N intake), and plasma Lys
concentration (Brito et al. 2009), and these positive
animal responses were likely attributed to greater microbial N production for cows fed high-WSC alfalfa (Brito
et al. 2009). Brito et al. (2009) also made several interesting observations; although cows fed high-WSC
alfalfa had greater DMI and total tract OM digestibility,
they tended to have higher rumen pH, which is consistent
with previous findings from studies evaluating effects of
sugar supplementation (Chamberlain et al. 1993; Heldt

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et al. 1999; Penner and Oba 2009). In addition, cows

fed high-WSC alfalfa increased BW to a lesser extent
compared with those fed low-WSC alfalfa. The effect of
feeding high-WSC forages on energy partitioning is of
interest for future research. However, high-WSC or highnon-structural-carbohydrate forages, which were evaluated in the past, also contained more starch and less
NDF compared with respective control forages. Therefore, the positive effects of high-sugar forages cannot
be solely attributed to specific effects of sugar.
Whey Products
Whey is a by-product of cheese manufacturing processes, and is available either as liquid or dried products.
Whey contains 70% lactose on a DM basis, and the
feeding value of whey products was extensively studied
in the United States in the late 1960s to early 1970s.
Research showed that feeding whey has similar effects
on rumen fermentation as feeding lactose, regardless of
whey type; butyrate concentration increased in lactating dairy cows (Huber et al. 1967), in sheep (Anderson
1975), and in dairy steers (Bragg et al. 1986). Rumen pH
was not affected by a partial replacement of corn grain
with dried whey (Huber et al. 1967; Bragg et al. 1986).
Huber et al. (1967) showed that milk production was not
affected by feeding dried whey in place of ground corn
and soybean meal at 30% of dietary DM or less, but
decreased at 60% of dietary DM. However, this response might be attributed to high dietary CP or ash
content of dried whey. Substituting dried whey for corn
grain and soybean meal at 30% of dietary DM increased
DMI, milk fat concentration, and 4% fat-corrected
milk yield without affecting milk yield (Casper and
Schingoethe 1986). More recently, Charbonneau et al.
(2006) showed that feeding dried whey permeate in place
of wheat starch or ground corn at 10.6% of dietary DM
increased DMI, but decreased milk yield. The possible
negative effects may be attributed to altered rumen
fermentation due to greater mineral intake and subsequent increase in water intake for cows fed whey products (Charbonneau et al. 2006). However, Anderson
et al. (1974) showed that average daily gain was increased for heifers fed grain and liquid whey compared
with those fed liquid whey or grain alone. Dried whey
can be fed to growing steers at 60% of dietary DM without causing digestive disorders or negatively affecting
growth (King and Schingoethe 1983). In an extensive
review, Schingoethe (1976) indicated that ruminants can
consume liquid whey up to 30% of dietary DM, and
feeding dried whey products more than 10% of dietary
DM in high grain diets may prevent milk fat depression.
Molasses
Molasses is another by-product feed with a high sugar
content. Broderick and Radloff (2004) conducted two
studies to determine the optimum dietary sugar content.
Increasing dietary sugar content up to 10% dietary DM
using dried or liquid molasses did not affect rumen pH or

volatile fatty acid profile, which agreed with findings from a recent study (Oelker et al. 2009). However,
ammonia concentration in rumen fluid was affected
quadratically (Broderick and Radloff 2004); the lowest
ammonia concentration was observed at dietary sugar
content of 4.2% in one study and 7.4% in the other
study, and diets containing the largest amount of sugar
in each study (7.2 and 10.0%, respectively) had greater
ammonia concentration. When dietary sugar content
was increased from 2.4 to 7.2% on a DM basis by a
partial replacement of high moisture corn with dried
molasses, DMI increased linearly, but milk yield and
composition were not affected. Thus, N utilization efficiency (milk N/N intake) was linearly decreased as
dietary sugar concentration increased, although urinary
N excretion decreased. When liquid molasses supplementation increased dietary sugar content up to 10.0%,
DMI increased quadratically, peaking at 4.9% of dietary
sugar concentration, but N utilization efficiency decreased linearly. It is also noteworthy that total tract
NDF digestibility was increased by molasses supplementation and peaked at the dietary sugar concentration of
7.2 or 7.4% (Broderick and Radloff 2004). This is not
consistent with in vitro studies that reported negative
effects of glucose addition on total tract NDF digestibility (Piwonka and Firkins 1993, 1996), but agrees with
the observations made for sucrose supplementation in
vivo (Broderick et al. 2008).
Other Feedstuffs
Citrus pulp is another high-sugar feedstuff. Lactating
dairy cows fed diets containing citrus pulp as a partial
replacement of corn grain increased the molar proportion of butyrate in the rumen fluid, but decreased
DMI, blood glucose concentration, and milk production
(Broderick et al. 2002). However, citrus pulp is rich in
pectin content, so the negative effects of feeding citrus
pulp cannot be solely attributed to greater dietary sugar
content, but also to greater pectin content or reduced
starch content.
Several commercial products that are high in sugar
content are also available. Firkins et al. (2008) reported
that feeding a liquid feed containing cane molasses and
corn steep liquor (Cargill Molasses Liquid Products
Division, Elk River, MN) tended to increase DMI, but
did not have consistent effects on milk yield and composition. Golombeski et al. (2006) evaluated a blend of
whey permeate and corn steep liquor as a source of
highly fermentable sugar, and reported that feeding the
highly fermentable sugar in place of corn grain and
soybean meal increased butyrate concentration in rumen
fluid and tended to increase milk yield, but tended to
decrease milk protein yield. However, corn steep liquor
has a high lactic acid content, which may confound the
effects of sugars on animal performance.

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OBA * FEEDING SUGARS AND DAIRY COWS 43

Associative Effects of Feeding Sugars

Synchronizing rates of carbohydrate fermentation and
protein degradation is, in theory, expected to increase
microbial protein production (Nocek and Russell 1988).
Kim et al. (1999a, b) conducted a series of studies to
evaluate how synchrony of release of energy and nitrogen in the rumen affects microbial protein production.
They infused sugar into the rumen continuously for 6 h,
beginning right after feeding (synchronized) or 6 h after
feeding (unsynchronized). Although the timing of ruminal sugar infusion did not affect microbial protein
synthesis in non-lactating cows fed grass silage only
(Kim et al. 1999a), lactating dairy cows consuming a
readily fermentable diet increased microbial protein
synthesis with the synchronized sugar infusion, but not
with the unsynchronized infusion (Kim et al. 1999b).
Similarly, ruminal infusion of sucrose increased urinary
excretion of allantoin, an indicator of microbial protein
production in the rumen, when infusion was conducted
at a time when expected availability of rumen degradable protein (RDP) was maximized (Trevaskis et al.
2001). These data support the concept that nutritional synchrony potentially increases microbial protein
production.
Sugars ferment faster than starch in the rumen, and it
was hypothesized that the positive effects of feeding
sugar would be greater for cows fed diets high in RDP
or soluble intake protein (SIP). However, this concept
has not been supported by data in the literature (Hall
and Huntington 2008). Casper and Schingoethe (1989)
evaluated three sources of carbohydrate varying in
fermentability (corn, barley, and whey) and two sources
of N differing in solubility (soybean meal and urea),
but did not observe animal responses supporting this
concept. Similarly, Sannes et al. (2002) reported that
feeding soybean meal or urea in high sugar diets did not
affect microbial protein synthesis. Dietary RDP level
(Arroquy et al. 2004a) or sources (Arroquy et al. 2004b)
did not affect nutrient digestibility of beef cattle fed diets
supplemented with dextrose or starch. Furthermore,
Hall et al. (2010) reported that the efficiency of fat- and
protein-corrected milk production was greater when
high-sugar feedstuffs (citrus pulp and molasses
sucrose) are fed in low RDP diets while the efficiency
was greater when ground corn was fed in a high RDP
diet. These results might be attributed to that sugarutilizing microbes preferentially use amino acids for
their growth rather than ammonia-N (Maeng and
Baldwin 1976). Although the importance of synchronizing energy and nitrogen availability was demonstrated
under the experimental settings (i.e., infusion), there is
no evidence in the literature demonstrating the need for
high-sugar diets to be fed with greater dietary RDP or
SIP.
CONCLUSIONS
Sugar ferments rapidly in the rumen, but feeding sugar
in place of dietary starch sources does not necessarily

decrease rumen pH. Thus, sugar may be used as an

alternative carbohydrate source in diets of lactating
dairy cows to increase the energy density of diets with
reduced risk of rumen acidosis. However, data in the
literature collectively indicate that feeding sugar may
not improve N utilization efficiency. Feeding high-sugar
diets is often associated with greater butyrate concentration in the rumen and is expected to enhance cell
proliferation, but its physiological mechanisms and the
effects of butyrate metabolism on the overall productivity of dairy cows warrant further investigation.
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