J. Dairy Sci.
85:1176–1182
 American Dairy Science Association, 2002.
Chewing Activity, Saliva Production, and Ruminal pH of Primiparous
and Multiparous Lactating Dairy Cows1
M. Maekawa,* K. A. Beauchemin,† and D. A. Christensen*
*Department of Animal and Poultry Science,
University of Saskatchewan,
Saskatoon, Canada, S7N 5A8
†Livestock Science Section,
Agriculture and Agri-Food Canada,
Research Centre, Lethbridge, Alberta, Canada, T1J 4B1
ABSTRACT
Four multiparous (MP) and four primiparous (PP)
ruminally cannulated lactating Holstein cows were
used in a double 4 × 4 Latin square design to study the
chewing behavior, saliva production, and ruminal pH
of cows in the first or subsequent lactation. Cows were
fed one of four diets; three total mixed rations con-
taining 40, 50, or 60% silage (DM basis), and a separate
ingredient diet containing 50% concentrate. Dry matter
intake was higher for MP cows than for PP cows (19.2
vs. 17.1 kg/d) but not as a percentage of body weight
(2.97 ± 0.06%). Multiparous cows spent more time
eating than PP cows (260 vs. 213 min/d, respectively),
even after adjustment for dry matter intake (13.8 vs.
12.4 min/kg DM). Multiparous cows also spent more
time ruminating per day than PP cows (560 vs. 508
min/d, respectively). Eating salivation rate was not af-
fected by parity, but resting salivation rate was higher
for MP cows than for PP. Although MP cows spent
more time chewing than PP cows, total daily saliva
production was only numerically higher for MP cows
because the increase in saliva produced during chewing
was accompanied by a decrease in saliva produced dur-
ing resting. Furthermore, pH profiles tended to be lower
for MP cows than for PP cows. Multiparous cows may
have a greater risk of incurring acidosis than PP cows
because increased salivary secretion associated with
increased chewing may not sufficiently compensate the
increment of fermentation acids produced in the rumen
due to high feed intake.
(Key words: parity, chewing activity, saliva pro-
duction)
Received March 5, 2001.
Accepted November 23, 2001.
Corresponding author: K. A. Beauchemin; e-mail: beauchemin@
em.agr.ca.
1
LRC contribution number 38701014.
1176
Abbreviation key: F:C = forage-to-concentrate ratio,
MP = multiparous cows, PP = primiparous cows, SI =
diet with ingredients allocated separately.
INTRODUCTION
Multiparous cows (MP) consume more feed per day
than do primiparous (PP) cows, and this is usually
reflected in higher milk production (Dado and Allen,
1994). Beauchemin and Rode (1994) suggested that
compared with MP cows, PP cows require longer eating
time and less competition at the manger to maximize
feed intake. Primiparous cows reportedly chew feed
more thoroughly and more slowly than older cows
(Campling and Morgan, 1981; Beauchemin and Rode,
1994). Thus, feed intake and chewing behavior differs
by parity (Grant and Albright, 1995).
Chewing activity is usually a good indication of ru-
men health because chewing stimulates saliva secre-
tion. Saliva has been estimated to supply about 70 to
90% of the fluid and buffering capacity entering the
rumen (Kay, 1966) and is the major determinant of
liquid outflow rate from the rumen (Cassida and Stokes,
1986). Silanikove and Tadmor (1989) reported a linear
relationship between feed intake and salivary secre-
tion rate.
Despite the importance of salivary secretion in the
digestive function of dairy cows, to our knowledge, no
studies have been conducted to quantify salivary secre-
tion during mastication as affected by parity. Further-
more, there is limited information in the literature re-
lated to differences in chewing activity and ruminal pH
of cows as affected by parity. Thus, the objective of this
experiment was to determine the effects of parity on
chewing activity, saliva production, and ruminal pH of
lactating dairy cows.
MATERIALS AND METHODS
Animals and Diets
Four MP and four PP ruminally cannulated Holstein
cows were used in this experiment. At the start of the
 CHEWING ACTIVITY, SALIVA PRODUCTION, AND PARITY
experiment, the MP cows averaged (± SD) 631 ± 87 kg
of BW and 109 ± 67 DIM. The PP group averaged 581
± 41 kg of BW and 146 ± 69 DIM. The experiment was
designed as a double 4 × 4 Latin square, with MP and
PP cows assigned to each square, respectively.
Cows received one of four diets. Three of the diets
were provided as a TMR consisting of different forage-
to-concentrate (F:C) ratios (40:60, 50:50, and 60:40),
whereas for the fourth diet, forage and concentrate were
fed separately (SI), and F:C ratio of the diet offered
was 50:50 (DM basis). The forage consisted entirely of
whole crop barley silage and the concentrates consisted
mainly of steam-rolled and ground barley grain. Fur-
ther information on diet composition and chemical com-
position of ingredients and diets is given in Maekawa
et al. (2002).
Measurements for the PP group were collected 7 d
apart from the MP group. All cows were cared for in
accordance with guidelines of the Canadian Council on
Animal Care (Ottawa, ON, Canada). Cows were housed
in individual stalls on rubber comfort mats with wood
shavings in the Dairy Unit of the Agriculture and Agri-
Food Canada Lethbridge Research Centre. Cows were
milked in their stalls twice daily. The cows were turned
outside for 1 or 2 h daily, except during measurements
of chewing activity, saliva secretion, and ruminal pH.
Cows were weighed at the beginning and end of each
period.
Feed Intake
The TMR was offered twice daily (0800, 1500 h) at
115% of voluntary intake. Forty percent of the daily
allocation was provided at the morning feeding, and
60% at the afternoon feeding. For cows fed SI, concen-
trate was fed at the same time as the TMR diets and
the barley silage was fed 1 h later (0900, 1600 h). Equal
proportions of concentrate were allocated at each meal,
but proportions of silage at each meal were the same
as for TMR. Orts for each cow were weighed daily.
Samples were taken daily during the last 6 d of the
period, composited by period for each cow, and dried at
55°C to a constant weight. Daily DMI was calculated
as the difference between the total amount of DM of-
fered and the DM refused, divided by 6 d.
Milk Yield and Components
Milk production was recorded daily at 0600 and 1630
h during the entire period. Milk samples were taken
a.m. and p.m. during the last 4 d of each period. They
were preserved with potassium dichromate, stored at
4°C, and sent to the Central Alberta Milk Testing Labo-
ratory (Edmonton, AB, Canada) for fat, protein, and
1177
lactose determination using an infrared analyzer (Milk-
0-Scan 605; Foss Electric, Hillerød, Denmark).
Chewing
Eating and ruminating behaviors were monitored vi-
sually for cows in both groups over a 24-h period. Eating
and ruminating activities were noted every 5 min, and
each activity was assumed to persist for the entire 5-
min interval. To estimate time spent eating or ruminat-
ing per kilogram of DM, NDF, or ADF intake, the aver-
age intake for the period was used. A period of rumina-
tion was defined as at least 5 min of ruminating activity
followed by at least 5 min without ruminating activity.
Total time spent chewing was calculated as the total
time spent eating and ruminating. Total time spent
resting was calculated as 24 h minus total time spent
chewing.
Salivation
The total amount of saliva secreted during eating
each day was calculated by multiplying the eating sali-
vation rate (ml/min) by the time spent eating each day
(min). Salivation rate during eating was measured by
collecting swallowed boluses of ingested feed from the
cardia during timed intervals as described in detail by
Maekawa et al. (2002). The amount of saliva added
to the masticate was calculated as the difference in
moisture of the collected bolus and the feed offered as
follows: saliva (ml) = weight of bolus (g) – weight of feed
as fed (g).
The total amount of saliva secreted during rumina-
tion each day was calculated similarly; however, the
rate of salivation during rumination was assumed to
be the same as that during eating (Bailey and Balch,
1961; Seth et al., 1974), because it was not possible to
measure this directly.
The total amount of saliva secreted during resting
each day was calculated by multiplying resting saliva-
tion rate (ml/min) by the time (min) spent resting each
day. Resting salivation rate was measured by collecting
swallowed saliva during timed intervals between feed-
ing times, as described in detail by Maekawa et al.
(2002). During this collection phase, feed was removed
to prevent eating, and water was turned off to prevent
cows from drinking. Volume of saliva was measured
immediately after each collection and resting salivation
rate was calculated by dividing the total amount of
saliva collected by the duration of each collection.
Ruminal pH
Ruminal pH was measured continuously for 24 h us-
ing an indwelling electrode, as described by Maekawa
Journal of Dairy Science Vol. 85, No. 5, 2002
1178 MAEKAWA ET AL.

Table 1. Milk production and composition.

Item Multiparous Primiparous SE P-value

DMI, kg 19.2 17.1 0.4 <0.01
BW, kg 639 590 5 <0.01
DMI, % of BW 3.03 2.90 0.06 0.15
Yield, kg/d
Milk 29.33 25.03 0.53 <0.01
4% FCM 28.01 24.21 0.54 <0.01
Fat 1.09 0.95 0.26 <0.01
Protein 0.95 0.83 0.02 <0.01
Lactose 1.35 1.19 0.03 <0.01
Composition, %
Fat 3.75 3.84 0.07 0.43
Protein 3.33 3.32 0.02 0.88
Lactose 4.56 4.76 0.02 <0.01

et al. (2002). Ruminal pH data were summarized for where

each cow in each period as daily mean pH, maximum
pH, minimum pH, minutes pH was below 5.8, and area µ =overall mean,
between the curve and pH 5.8. The area was calculated Si =random effect of parity (i = 1 to 2),
by adding the absolute value of negative deviations in Ln =effect of stage of lactation (n = 1 to 3),
pH from pH 5.8 for each 15-min interval. Data from (SL)in =interaction of parity and stage of lactation,
one cow in period 4 was not used due to problems with and
the pH electrode. ein = residual.

Rate of Passage of Rumen Liquid Fraction
The rate of passage of the liquid fraction from the
rumen was measured using Co-EDTA, as described by
Maekawa et al. (2002). The last day of each period,
rumen contents were completely removed and weighed.
Samples were taken for DM analysis to calculate rumen
liquid volume. The digesta from each cow were switched
to the cow receiving that diet the next period to mini-
mize the need for a lengthy adaptation period.
Significance was declared at P < 0.05 and trends are
discussed at P < 0.15. Because there were no interac-
tions between diet and parity of cows, only the least
squared means for parity are presented. Effects of di-
etary treatments were reported separately (Maekawa
et al., 2002).
RESULTS
DMI, BW, Milk Yield, and Composition

Statistical Analyses
For each period, means for individual cows were cal-
culated for all variables. Data were analyzed as a double
4 x 4 Latin square using the general linear model proce-
dure in SAS (1989) to account for the effect of parity,
cow within parity, period within parity, diet, and the
interaction between parity and diet. Because there were
no interactions between diet and parity of cows, only
the effects of parity are reported. The effect of diet is
reported separately (Maekawa et al., 2002).
To analyze the effect of stage of lactation on DMI and
eating saliva, cow and period effects were eliminated
from the analyses, and the data were analyzed as a
randomized complete block design, using the general
linear model procedure of SAS (1989). The effect of block
was assumed to be the effect of parity. The model
used was:
Yij = µ + Si + Ln + (SL)in + ein
Dry matter intake was higher for MP cows than for
PP cows, but MP cows were heavier than PP cows (Table
1). Higher DMI of MP cows reflected the heavier BW of
the MP cows, because there was no difference between
groups when DMI was expressed as percentage of BW.
Multiparous cows produced 4.3 kg/d (17%) more milk
than PP cows (Table 1). Even after correction for fat
content, MP cows produced 17% more milk than PP
cows. Yield of fat, protein, and lactose were also higher
for MP cows compared with PP cows. However, differ-
ences in fat and protein yields between parity groups
were due to differences in milk yield, not differences
in component percentages. Multiparous cows produced
milk with lower lactose percentage than PP cows, and,
consequently, lactose yield was lower for MP cows.
Chewing Activity
Multiparous cows spent 47 min/d longer eating than
PP cows (Table 2). Multiparous cows also tended (P =

Journal of Dairy Science Vol. 85, No. 5, 2002

 CHEWING ACTIVITY, SALIVA PRODUCTION, AND PARITY 1179
Table 2. Chewing activity of multiparous and primiparous lactating dairy cows.

Item Multiparous Primiparous SE P-value

Eating
Min/d 260 213 8 <0.01
Min/kg of DM 13.8 12.4 0.5 0.07
Rumination
Min/d 560 508 17 0.05
Min/kg of DM 29.3 29.6 0.9 0.78
Min/kg of ADF 163 164 5 0.81
Min/kg of NDF 99 100 3 0.80
Numbers of periods 14.1 15.9 0.6 0.05
Periods/kg of DM 0.75 0.94 0.03 <0.01
Length of periods, min 40.7 32.7 1.7 0.01
Total chewing, min/d 821 720 19 <0.01
Resting
Min/d 619 720 20 <0.01

0.07) to spend 1.4 min more eating per kilogram of DM
than PP cows. Rumination time was also greater for
MP cows than for PP cows. Multiparous cows spent
52 min/d longer ruminating than PP cows. However,
increased rumination time likely reflected the higher
DMI of MP cows, because time spent ruminating per
kilogram of DM, ADF, or NDF intake was similar be-
tween groups. Despite longer rumination time per day,
MP cows had fewer rumination periods than PP cows.
Rumination periods per kilgram of DM were fewer for
MP cows compared with PP cows. However, MP cows
ruminated 8 min longer each period compared with
PP cows.
As a result of longer eating time and longer rumina-
tion time, MP cows spent more than 1.5 h longer each
day chewing (Table 2). Consequently, MP cows spent
less time resting compared with PP cows.
Saliva Production
Salivation rate during eating was similar for MP and
PP cows (Table 3). However, salivation rate during rest-
ing tended (P = 0.06) to be higher for MP cows than for
PP cows. Daily total secretions of saliva during resting
and rumination were similar for both groups. However,
MP cows secreted more saliva per day during eating
than did PP cows. However, because the quantity of
saliva secreted during eating represented only approxi-
mately 22% of the total daily saliva production, MP
cows had only numerically higher total saliva secretion
per day than PP cows (P = 0.16).
Ruminal pH
The ruminal pH variables measured were similar for
both groups, even though mean pH was 0.11 units lower
for MP cows than for PP cows (Table 4). Multiparous
cows also remained under pH 5.8 about 1.2 h/d longer
(P = 0.25) and minimum pH was 0.05 units lower (P =
0.39) than for PP cows.
The pattern of diurnal fluctuation of ruminal pH was
similar between parities (P = 0.10) (Figure 1). Ruminal
pH decreased immediately after the 1500 h meal, then
increased during the night and decreased again after
the morning meal. Multiparous cows showed greater
decline in ruminal pH than PP cows after the afternoon
meal, although mean pH from 1500 to 0245 h was simi-
lar for both groups (Table 4). However, mean pH calcu-
lated for the period of 0300 to 1445 h was lower for MP
cows than for PP cows.

Table 3. Saliva production during eating and resting for multiparous and primiparous lactating cows.
Item Multiparous Primiparous SE P-value
Eating
Salivation rate, ml/min 225 226 11 0.93
Resting
Salivation rate, ml/min 114 88 9 0.06
Saliva output, L/d
Eating 56 49 2 0.02
Resting 70 63 6 0.36
Ruminating1 125 115 8 0.44
Total 252 227 12 0.16
1
Rate of salivary secretion was assumed to be the same during ruminating as during eating.

Journal of Dairy Science Vol. 85, No. 5, 2002

1180 MAEKAWA ET AL.

Table 4. Ruminal pH of multiparous and primiparous lactating dairy cows.

Item Multiparous Primiparous SE P-value

Mean pH 5.84 5.95 0.06 0.25
Minimum pH 5.25 5.30 0.04 0.39
Maximum pH 6.54 6.56 0.05 0.77
Area under pH 5.8, pH × h/d 3.54 3.20 0.57 0.71
Time under pH 5.8, h/d 11.42 8.93 1.33 0.25
Percent of time under pH 5.8, % 47.63 37.18 5.59 0.25
Mean pH from 15:00 to 02:45 5.74 5.75 0.07 0.97
Mean pH from 03:00 to 14:45 5.94 6.15 0.06 0.05

Rumen Liquid Turnover
The MP cows had higher rumen digesta weight than
the PP cows, due to the higher liquid content because
the amount of DM in the rumen was similar for both
groups (Table 5). The MP cows had higher liquid outflow
rate than PP cows and they also had higher liquid turn-
over rate. Higher liquid content in the rumen of MP
cows compared with PP cows was associated with larger
rumen volume due to body size; as the relation between
rumen liquid volume and BW was similar for both
groups.
DISCUSSION
Dry matter intake was positively correlated with BW,
thus MP, larger cows ate more than PP, smaller cows
as reported previously (Kertz et al., 1991; Beauchemin
and Rode, 1994; Dado and Allen, 1994). Milk production
was also positively correlated with DMI, accounting for
the higher milk and FCM yield of MP cows than PP
cows, as reported previously by others (Colenbrander
et al., 1991; Dado and Allen, 1994; Beauchemin et al.,
1997). However, effects of parity on milk composition
are less clear. Colenbrander et al. (1991) fed cows alfalfa
silage and corn grain and observed no difference in
milk fat percentage between MP and PP cows, as was
observed in the present study. However, when cows
were fed alfalfa hay and barley concentrate, Beauche-
min and Rode (1994) found that milk of MP cows con-
tained similar fat content but less protein and less lac-
tose than the milk of PP cows. In another experiment,
Beauchemin and Rode (1997) fed diets consisting of
three percentages of NDF from barley silage combined
with concentrates based on either barley or corn, and
found that MP cows yielded more milk, of lower fat
content, than did PP cows. This, milk composition is
likely influenced more by factors such as stage of lacta-
tion of the cows, physical characteristics of the diet,
and proportion of fiber in the diet (Robinson, 1989; Tess-
man et al., 1991; Coulon et al., 1994) than by parity.
Chewing behavior was affected by parity, with higher
eating and ruminating activity for MP cows than for
PP cows. Previously, Beauchemin et al. (1997) found
that MP cows spent more time eating than PP cows,
although rumination time was similar for both groups.
In contrast, Beauchemin and Rode (1997) reported that
MP cows spent less time eating and more time ruminat-
ing than did PP cows, although MP cows spent less time
eating and ruminating per kilogram of DM or NDF. The
relationship between parity and time spent eating and
ruminating may depend on the type of diet. With high
fiber diets, cattle with a greater intake capacity (MP
cows) tend to chew feed more efficiently during eating
and ruminating, requiring less chewing time per unit
of feed DM consumed, because the relationship between
rumination capacity and body size is near unity (De
Boever et al., 1990). This was not the case in the present
study; MP cows did not chew more efficiently than
PP cows.
Beauchemin (1991) reported that the amount of sa-
liva secreted daily was proportional to feed intake. In
the present experiment, there was also a significant
relationship between DMI and the total amount of sa-
liva produced during eating (r = 0.58, P = < 0.01; Figure

Table 5. Passage rate of the rumen liquid fraction for multiparous and primiparous lactating dairy cows.

Item Multiparous Primiparous SE P-value

Rumen digesta weight, kg 93 86 2 0.05
Rumen content, kg of DM 13.3 13.2 0.4 0.71
RLV1, L 79 73 2 0.03
Liquid outflow, L/h 9.52 7.99 0.30 <0.01
Liquid turnover rate, %/h 12.04 11.13 0.22 0.01
RLV/BW, L/kg 0.125 0.124 0.002 0.85
1
RLV = Rumen liquid volume.

Journal of Dairy Science Vol. 85, No. 5, 2002

 CHEWING ACTIVITY, SALIVA PRODUCTION, AND PARITY
Figure 1. Diurnal fluctuation in ruminal pH of multiparous
(dashed) and primiparous (solid line) lactating dairy cows. Each line
represents the mean of 16 observations. Measurements were collected
every 15 s and averaged for each 15-min interval. Arrows indicate
feeding time.
2). Thus, higher salivary secretion during eating by
MP cows compared with that of PP cows was partly in
response to higher DMI. Cassida and Stokes (1986)
reported that stage of lactation influenced saliva pro-
duction, with reduced saliva production for cows in
early lactation. However, stage of lactation also influ-
ences DMI. In the present study, stage of lactation in-
fluenced salivary secretion during eating to a lesser
extent than DMI (Figure 3). The decrease in DMI for
cows in late lactation was not accompanied by a de-
crease in salivary secretion during eating.
Although salivation rate during resting was higher
for MP cows, there was no affect of parity on total
amount of saliva secreted during resting because MP
cows spent less time resting than PP cows. The higher
salivation rate during resting for MP cows may be an
involuntary response to reduced time spent resting and
the necessity to buffer the high level of fermentation
acids. Although MP cows ruminated 52 min/d longer
than PP cows, the net increase in saliva production was
only about 5 L/d. This is because the 10-L increase in
saliva output during rumination was accompanied by
a 5-L decrease in saliva output during resting (Table 3).
Even though only numerical differences between MP
and PP cows were observed for the ruminal pH vari-
ables measured, these differences are likely to be of
biological significance in terms of lowering fiber diges-
tion. A larger study with a greater number of cows is
needed to confirm the trend of lower ruminal pH for
MP cows.
Higher rumen liquid volume of MP cows compared
with PP cows could be the result of higher production
of saliva and higher water intake. Although in our ex-
periment water intake was not measured, Dado and
Allen (1994) found that MP cows drank 30% more water
than PP cows. Increased liquid outflow rates, as was
observed for MP cows compared with PP cows is thought
to be beneficial, in terms of increased delivery of metab-
1181
Figure 2. Relationship between DMI and saliva secreted during
eating. Eating saliva (L/d) = −0.51 + 2.59 ( DMI (kg/d), r = 0.58, P =
< 0.01.
olizable protein to the lower gut (Russell and Hes-
pell, 1981).
CONCLUSION
Salivation rate during eating was not affected by par-
ity, but the greater eating time of MP cows resulted in
greater saliva output during eating. However, saliva
produced during eating only accounted for 22% of the
total daily saliva output. Salivation rate during resting
was higher for MP cows than for PP, which may be an
involuntary response to reduced time spent resting and
the necessity to buffer the rumen environment. Al-
though MP cows ruminated more than PP cows, the
net increase in saliva production was small, because of
the accompanying decrease in resting saliva. Thus,
daily saliva production was only numerically higher
for MP cows. Furthermore, production of fermentation
acids in the rumen was likely higher for MP cows due
to higher feed intake, thus pH profiles of MP cows
tended to be lower than for PP cows. Multiparous cows
may have a greater risk of incurring acidosis than PP
Figure 3. Relationship between saliva secreted during eating
(䊐)(SE = 4.66, 2.57, and 3.20, for early, medium, and late lactation,
respectively) and DMI (SE = 1.01, 0.56, and 0.69, for early, medium
and late lactation, respectively) and stage of lactation of dairy cows.
Each point represents the mean of 5, 16, and 11 observations, for
early, medium, and late lactation, respectively.
Journal of Dairy Science Vol. 85, No. 5, 2002
1182 MAEKAWA ET AL.
cows because the increased salivary secretion associ-
ated with increased chewing may not sufficiently com-
pensate for the higher production of fermentation acids
in the rumen due to higher feed intake.
ACKNOWLEDGMENTS
The authors wish to thank the staff of the Lethbridge
Research Centre Dairy Unit for caring for the animals
and G. Bowman, S. Eivemark, B. I. Farr, J. Bovinec,
and J. Chang for their technical assistance during ex-
perimentation. The authors also thank to W. Yang and
K. Koenig for reviewing this manuscript. Agriculture
and Agri-Food Canada and the Canada/Alberta Live-
stock Research Trust Inc. funded the project.
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