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Rumen Fermentation and Nutrient Flows for Cows

Fed Grass and Grass Supplemented


with Molassed Beet Pulp Pellets
F. P. O’MARA,*,1 G. K. STAKELUM,* P. DILLON,*
J. J. MURPHY,* and M. RATH†
*Teagasc, Dairy Husbandry Department, Moorepark,
Fermoy, Co. Cork, Ireland
†Department of Animal Science and Production,
University College Dublin, Belfield, Dublin 4, Ireland

ABSTRACT INTRODUCTION
An experiment was carried out to determine the Rumen digestion is characterized by the degrada-
effect of a grass diet and a concentrate supplement on tion and fermentation of ingested carbohydrate and
rumen fermentation and nutrient flows to the duode- protein in the rumen. By-products arising from this
num. Perennial ryegrass was cut and fed indoors to process are absorbed or resynthesized into microbial
eight rumen- and duodenum-cannulated Friesian biomass. Therefore, a substantial quantity of the
cows with or without 3 kg/d of molassed beet pulp in a nutrients that are available for absorption is of
randomized design experiment. The dry matter in- microbial origin. To understand the responses of
take of grass was significantly lower for cows fed the ruminants to different feedstuffs, it is vital to know
concentrate supplement (13.6 vs. 11.5 kg of dry which nutrients are absorbed by ruminants. Thus,
much effort has been expended on measuring rumen
matter/d), but total dry matter and organic matter
fermentation and nutrient flows to the duodenum in
( O M ) intakes were similar for cows fed both diets.
ruminants. This research has generally involved diets
Cows fed the supplement had higher mean concentra-
in which large amounts of concentrates are fed with
tions of total volatile fatty acids (108 vs. 89 mmol/L) conserved forages. However, in many temperate
and a higher percentage of butyrate in total volatile regions, the bulk of ruminant diets is grass obtained
fatty acids (13.5 vs. 11.6 mol/100 mol). There were by grazing. Performance of cows fed fresh grass diets
no differences between the diets in the flow of OM to can surpass that of cows fed silage conserved from the
the duodenum or in the extent of OM digestion in the same grass ( 1 7 ) and is often better than the perfor-
rumen. Flows of nonammonia N, microbial N, and mance of cows fed conserved silage supplemented
amino acids to the duodenum tended to be higher for with moderate amounts of concentrates. For example,
cows fed the supplemented diet than for those fed higher milk production has been reported for cows
ryegrass only. The efficiency of microbial protein syn- that grazed on grass throughout most of their lacta-
thesis also tended to be higher for cows fed the sup- tion than for cows that were fed a higher percentage
plemented diet (42 vs. 37.7 g/kg of OM apparently of grass silage and concentrates in the diet (12).
digested in the rumen and 28.2 vs. 26 g/kg of OM Despite the importance of grass as a feed in tem-
truly digested in the rumen). Overall, there were perate regions, there are few reports of nutrient flows
indications that the supplement caused better capture to the duodenum of dairy cows when diets consisted
of N in the rumen and increased the efficiency of predominantly of fresh grass, and none of these
microbial protein synthesis. studies examined the effect of the supplementation of
( Key words: dairy cows, grass, beet pulp, digestion) concentrates. The objective of this experiment was to
provide such information for lactating dairy cows that
Abbreviation key: DOMI = digestible OM intake, were fed diets of fresh grass and to determine the
OMAD = OM apparently digested in the rumen, effect of a supplement of molassed beet pulp on rumen
OMD = OM digestibility, OMTD = OM truly digested fermentation and nutrient flows to the duodenum.
in the rumen, SED = standard error of the difference.
MATERIALS AND METHODS

Received May 10, 1996. Experimental Diets and Treatments


Accepted April 22, 1997.
1Current address: Department of Animal Science and Produc- Eight multiparous Friesian cows were allocated at
tion, University College Dublin, Belfield, Dublin 4, Ireland. random to one of two dietary treatments: grass only

1997 J Dairy Sci 80:2466–2474 2466


NUTRIENT FLOWS FOR COWS FED GRASS DIETS 2467

or grass supplemented with 3 kg/d of molassed beet duodenum (14). Beginning on d 19 of the experiment,
pulp pellets (International feed identification no. approximately 29.73 Ci/d of 51Cr and 2.16 Ci/d of
103Ru in 1 L of distilled water were continuously
4-00-672). Cows fed the unsupplemented and sup-
plemented diets averaged 68 and 70 DIM, respec- infused into the rumen of each cow using multichan-
tively. The grass offered was predominantly perennial nel peristaltic pumps. The volumes of liquid infused
ryegrass, which was cut once daily with a reciprocat- daily were recorded to determine the exact amounts
ing mower type grass cutter (Vakuumat, Wartberg, of the markers that were infused.
Austria) at a height of 5 to 6 cm. Cutting started on On d 25 of the experiment, duodenal contents were
April 16th and continued for 40 d. For the first 20 d, sampled four times at 0600, 1200, 1800, and 2400 h.
When sampling, the first pulse was discarded, and
the grass cut was primary growth. Beginning on d 21,
approximately 500 ml were taken at each sampling
the grass was cut from an area in which the primary
time. Composite samples of whole duodenal contents
growth had been cut 28 d previously. The cows were and the residues after centrifugation at 1000 × g for
tied in individual stalls and were fed and milked in 10 min at 0°C were prepared for each cow and stored
these stalls. Cows were milked at 0830 and 1630 h. at –20°C. Bacterial pellets were prepared for each cow
Milk production was recorded daily, and milk compo- by centrifuging the supernatant at 6000 × g for 30
sition was determined on 1 d/wk. min at 0°C. The pellets were washed in distilled
water and centrifuged again at 6000 × g. Pellets were
Management and Procedures stored at –20°C and freeze-dried before chemical
analysis.
Herbage was fed six times each day at 1000, 1200, The samples of duodenal contents (whole and
1400, 1700, 2000, and 2400 h. The beet pulp supple- residue after centrifugation) were composited and
ment was fed once daily at approximately 0930 h. For milled through a 1-mm screen after freeze-drying.
the first 17 d, herbage was fed for ad libitum intake Ash and CP concentrations were determined as
while cows adapted to the diets. Beginning on d 18, described previously. The amino acid content of these
herbage allowance was restricted to 95% of mean composite samples was determined by ion-exchange
herbage DMI on d 15, 16, and 17. chromatography ( 1 9 ) using a Beckman analyzer
The amounts of herbage offered and orts were sam- (model 6300; Beckman Instruments, Fullerton, CA).
Methionine and cysteine were determined as meth-
pled every day, and weights were recorded. Dry mat-
sulfone and cysteic acid, respectively, on preoxidized
ter content was measured by drying at 95°C for 16 h
samples (19). Samples were analyzed for NH3 con-
in a forced-air oven. The dried samples were milled centration ( 2 ) . Concentrations of 51Cr and 103Ru
through a 1-mm screen, and ash content was deter- were analyzed simultaneously on a High Purity Ger-
mined by burning in a muffle furnace at 550°C for 16 manium P type detector (Canbera-Packard Instru-
h. To calculate feed allowances, a rapid estimation of ments Ltd., Pangbourne, Berkshire, UK) connected to
DM was also carried out each day; four 50-g subsam- a Canbera series 20 multichannel analyzer. Concen-
ples were dried at 130°C for 30 min in a forced-air trations of 51Cr and 103Ru were used to determine
oven. Fresh herbage samples were frozen at –20°C, flow rates (14). Purine concentrations in digesta
freeze-dried, and ground through a 1-mm screen be- samples and bacterial pellets, which had been ground
fore analysis of Kjeldahl CP concentration ( 2 ) using with a pestle and mortar, were measured ( 3 3 ) to
a Kjelfoss instrument (Foss Electric, Hillerød, Den- determine microbial protein flow.
mark), modified ADF concentration (11), NDF con- Samples of rumen fluid (approximately 300 ml)
centration (29), and in vitro OM digestibility were taken through the rumen cannula at 0800, 0930,
( OMD) (21). 1100, 1500, 2000, and 2400 h on d 25 of the experi-
The molassed beet pulp pellets were sampled every ment. These samples were strained through four lay-
ers of cheesecloth, and then the pH was measured. A
day, composited by week, and milled through a
small sample ( 5 ml approximately) was acidified
1-mm screen. Dry matter content was determined by
with concentrated H2SO4 and stored at –20°C before
drying at 103°C for 4 h in a forced-air oven. Ash and subsequent analysis. The NH3 concentration was
CP concentrations were determined as described measured enzymatically using a Cobas Mira biochem-
previously. Weende crude fiber was also determined ical analyzer (Roche Diagnostica, Basle, Switzer-
(2). land). Volatile fatty acids were measured using an
The 51Cr-labeled EDTA and 103Ru-labeled Tris- automated gas chromatograph (Perkin Elmer Sigma
(1.10 phenantroline)-ruthenium (11)-chloride com- 3; Perkin Elmer Corp., Norwalk, CT) fitted with a
plexes were used as the liquid and solid phase mark- Tween 80 and Chromasorb WAW 80/100 column (Su-
ers, respectively, to determine nutrient flow to the pelco, Inc., Bellefonte, PA). D- and L-Lactate were
Journal of Dairy Science Vol. 80, No. 10, 1997
2468 O’MARA ET AL.

measured enzymatically using an express clinical TABLE 1. Chemical composition of the grass fed during the infu-
sion and sampling period and the fecal collection period.
chemistry analyzer (model 530; Ciba Corning Diag-
nostics, Oberlin, OH). Period
On d 30 of the experiment, tubes were fitted to Infusion Fecal
divert urine, and total feces output was measured for and sampling collection
the next 10 d to determine in vivo digestibilities. The DM, % 15.7 18.7
feces were sampled each day, dried at 60°C for 72 h, CP, % of DM 15.1 15.1
allowed to cool, weighed, ground through a 1-mm Ash, % of DM 8.0 8.0
Modified ADF, % of DM 21.1 23.0
screen, and dried at 103°C for 4 h before DM was NDF, % of DM 39.9 44.0
determined. Ash was determined as described previ- In vitro OM digestibility, % 81.8 79.1
ously, and DM digestibility and OMD were calcu-
lated.

Statistical Methods tions. The grass that was primary growth and that
The difference between the data for diets in feed was fed during the first 20 d of the experiment con-
intake, digestibility, rumen fermentation, and sisted of 18.6% modified ADF, 36.2% NDF, 18.2% CP,
nutrient flow were analyzed for a completely ran- and 8.5% ash in the DM; the in vitro OMD was 84.9%.
domized design by the ANOVA procedure of SAS The molassed beet pulp consisted of 89.6% DM, and
(24). The difference between the diets in rumen 7.7% ash, 12.1% CP, and 13.7% crude fiber in the DM.
parameters at each sampling time and for the calcu- Intakes of grass and beet pulp DM, OM, and N
lated mean value were analyzed by the ANOVA during the infusion and sampling period and the fecal
procedure of SAS. For all analyses, significance was collection period are presented in Table 2. Grass DMI
declared at P = 0.05. was lower ( P < 0.05) for cows fed the supplemented
diet during both periods. There was little difference in
RESULTS total DMI and OM intake between the diets during
either period, although both tended to be higher for
The milk yield during the course of the experiment cows fed the supplemented diet. Nitrogen intakes
was 21.4 and 19.8 kg/d [standard error of the differ- were similar for cows fed either diet. There was no
ence ( SED) = 3.83 kg/d] for cows fed the unsup-
plemented and supplemented diets, respectively. Milk
fat and protein concentrations were 4.24 and 4.27%
(SED = 0.223%) and 3.17 and 3.25% (SED = 0.208%)
for cows fed the unsupplemented and supplemented
diets, respectively. The mean live weight of the cows
during the experiment was 500 and 486 kg (SED =
35.9 kg) for cows fed the unsupplemented and sup-
plemented diets, respectively.
The chemical analysis of the grass offered to the
cows is presented in Table 1. Because of the possibil-
ity of changes in the composition of the grass over
time, the analysis is shown separately for the
11-d infusion and sampling period and the 10-d fecal
collection period (Table 1). The grass from both peri-
ods was similar in composition, except for slightly
higher modified ADF and NDF contents and lower in
vitro OMD values in the later cut grass, indicating
some maturation. Figure 1 shows that the in vitro
OMD was approximately 80.5% and was reasonably
constant throughout these two periods. The fiber con-
centrations indicate that the grass was composed of
leafy material, but the CP concentrations were not as
high as the primary spring growth. Grass in both
Figure 1. In vitro OM digestibility (OMD) of grass fed to cows
periods was cleanly cut and showed no evidence of soil in two dietary groups each day including the infusion and sampling
contamination as determined by the ash concentra- period ( d 19 to 25) and the fecal collection period ( d 30 to 40).

Journal of Dairy Science Vol. 80, No. 10, 1997


NUTRIENT FLOWS FOR COWS FED GRASS DIETS 2469

significant difference between the in vivo DM digesti- the two treatments, but OM truly digested in the
bility of the diets (80.4 and 80.9% for cows fed the rumen ( OMTD) averaged 74.9% of OM intake. The
unsupplemented and supplemented diets, respec- OMTD was calculated by adding the duodenal flow of
tively; SED = 0.37%), but the in vivo OMD of cows fed microbial OM to OMAD, assuming that the ash con-
the supplemented diet was higher ( P < 0.05) than tent of microbial DM was 10%.
that of cows fed the unsupplemented diet (82.2 and Duodenal flows of N and the various nitrogenous
81.1%, respectively; SED = 0.42%). components are presented in Table 5. As outlined in
Total VFA concentrations and the molar percent- Table 2, N intake was similar for cows fed either diet.
age of butyrate were higher ( P < 0.05) for cows fed There were no significant differences between the
the supplemented diet than for cows fed the unsup- diets in flows of total N or NAN, but flows were 31
plemented diet (Table 3). There were no other sig- and 32 g/d higher, respectively, for cows fed the sup-
nificant differences between the diets in the mean plemented diet than for cows fed the unsupplemented
values of any of the rumen parameters that were diet. Flow of NAN is the sum of microbial N and non-
measured. Changes in rumen pH, NH3, and total NH3, nonmicrobial N. The numerically higher NAN
lactate concentrations throughout the day are shown
flow for cows fed the supplemented diet was due to
in Figure 2. Rumen pH dropped more rapidly and
the microbial N flow, which tended to be higher (39 g/
reached lower levels (pH < 6 ) for cows fed the sup-
d ) for cows fed this diet. However, the difference in
plemented diet than for cows fed the unsupplemented
microbial N flow between the diets did not reach
diet. The difference in pH between treatments was
statistical significance. Microbial protein synthesis is
significant ( P < 0.05) at the third sampling time,
related to the amount of carbohydrate fermented in
1100 h, which was 90 min after the supplement was
the rumen and the yield of microbial N per kilogram
fed. The NH3 concentrations were higher at most
sampling times for cows fed the unsupplemented diet of carbohydrate fermented in the rumen. Carbohy-
than for cows fed the supplemented diet, but the drates fermented in the rumen were not determined
differences never reached statistical significance. The in this experiment, but microbial N flow per kilogram
pattern of change in total lactate concentrations of OMAD or per kilogram of OMTD tended to be
differed between the treatments, but the differences higher for cows fed the supplemented diet.
never reached statistical significance at any sampling There were no significant differences between the
time. diets in flows of total or individual AA (Table 6).
There were no significant differences between the Amino acid N flow, as a percentage of total N flow,
diets in duodenal flows of DM, OM, and OMD in the averaged 73.8%. The AA profile (grams of AA per 100
rumen (Table 4). Organic matter apparently digested g of AA) is presented in Table 7. The AA profile was
in the rumen ( OMAD) was calculated as the differ- similar for both diets, and the only significant differ-
ence between OM intake and OM flow at the duode- ence was a slightly higher percentage of Phe for cows
num. The OMAD averaged 51.4% of OM intake for fed the unsupplemented diet ( P < 0.05).

TABLE 2. Dry matter, OM, and N intakes during the infusion and sampling period and the fecal
collection period for cows fed unsupplemented grass diet ( U ) or grass diet supplemented with
molassed beet pulp pellets ( S ) .
Period
Infusion and sampling Fecal collection
U S P SED1 U S P SED
(kg/d) (kg/d)
Grass DMI 13.6 11.5 * 0.58 13.3 10.8 * 0.87
Pulp DMI 2.7 2.7
Total DMI 13.6 14.2 NS2 0.58 13.3 13.5 NS 0.87
Total OM intake 12.6 13.1 NS 0.54 12.3 12.5 NS 0.81
Total N intake 0.330 0.326 NS 0.0144 0.320 0.307 NS 0.021
1Standard error of the difference.
2P > 0.10.
*P < 0.05.

Journal of Dairy Science Vol. 80, No. 10, 1997


2470 O’MARA ET AL.

DISCUSSION TABLE 4. Duodenal flows of DM and OM from the rumen and


digestibility of OM in the rumen for cows fed unsupplemented grass
diet ( U ) or grass diet supplemented with molassed beet pulp
Intake pellets ( S ) .
Treatment
Herbage intake was depressed by 0.79 and 0.91 kg
of DM/kg of supplement during the infusion and sam- U S P SED1
pling period and the feces collection period, respec- Duodenal flow
tively. This high substitution rate agrees with the DM, kg/d 9.6 8.9 NS2 1.17
OM, kg/d 6.2 6.3 NS 0.49
results of previous researchers (8, 9, 15, 26) who OMAD3 6.4 6.8 NS 0.47
reported intakes in situations in which fresh cut grass OMAD, % of OM Intake 51.2 51.6 NS 3.07
was fed. Such a high substitution rate when grass is OMTD4 9.2 10.0 NS 0.52
OMTD, % of OM Intake 73.5 76.2 NS 2.38
fed for ad libitum intake indicates the difficulties of
trying to increase the DMI of grazing animals when 1Standard error of the difference.
grass supply is adequate. 2P> 0.10.
3OM apparently digested in the rumen.
4OM truly digested in the rumen.
Rumen Fermentation
The more rapid drop in pH for cows fed the sup-
plemented diet (Figure 2 ) indicated that the supple- ryegrass regrowth ( 3 ) , which is similar to the concen-
ment was fermented quite rapidly in the rumen. The tration that was detected for cows fed the unsup-
molasses proportion of the molassed beet pulp proba- plemented grass diet in this study (4.2 mmol/L).
bly underwent rapid fermentation. The pH values Rumen sampling took place in early to mid May in
remained >6 at most sampling times for both treat- this experiment, but the grass fed at this stage was
ments, which suggests that the supplement had no regrowth material.
detrimental effect on the activity of celluloytic bac- Supplements of molassed or unmolassed beet pulp
teria. have previously been reported to reduce rumen NH3
concentrations (25). The NH3 concentrations in this
Rumen degradation of herbage N is high ( 1 0 ) and
experiment followed the same direction, but the
can reach 80% or more of ingested N. High concentra-
difference was not significant. In another report (31),
tions of rumen NH3 are therefore possible for cows fed
grass diets, and mean daily values above 10 mmol/L
have been reported for cows consuming diets consist-
ing of primary growth of perennial ryegrass (5, 32) or TABLE 5. Flow of nitrogenous components at the duodenum for
late season perennial ryegrass regrowth ( 3 ) . Lower cows fed unsupplemented grass diet ( U ) or grass diet sup-
values (3.9 and 4.1 mmol/L) have been reported for plemented with molassed beet pulp pellets ( S ) .
diets consisting of midseason (July) perennial Treatment
U S P SED1
Total N
g/d 379 410 NS2 34.1
TABLE 3. Rumen fermentation data averaged over the six sam- % of N Intake 114 126 NS 8.8
pling times for cows fed unsupplemented grass diet ( U ) or grass NH3 N, g/d 22 21 NS 2.6
diet supplemented with molassed beet pulp pellets ( S ) . NAN Flow
Treatment g/d 357 389 NS 32.1
% of N Intake 108 119 NS 8.2
U S P SED1 NANMN3 Flow
g/d 116 109 NS 19.3
pH 6.7 6.5 NS2 0.25 % of N Intake 35 34 NS 5.9
NH3, mmol/L 4.2 3.3 NS 1.04 % of NAN 33 28 NS 4.6
D-Lactate, mmol/L 1.1 1.1 NS 0.33 Microbial N flow
L-Lactate, mmol/L 0.4 0.5 NS 0.09 g/d 241 280 NS 26.7
Total VFA, mmol/L 89 108 * 7.6 % of NAN 67 72 NS 4.6
Acetic acid, mol/100 mol 62.8 61.2 NS 1.14 g/kg of OMAD4 37.7 42.0 NS 5.41
Propionic acid, mol/100 mol 24.9 23.9 NS 0.92 g/kg of OMTD5 26.0 28.2 NS 2.55
Butyric acid, mol/100 mol 11.6 13.5 * 0.61
Isobutyric acid, mol/100 mol 0.8 0.9 NS 0.58 1Standard error of the difference.
C2:C3 2.6 2.6 NS 0.12 2P > 0.10.
1Standard error of the difference. 3Non-NH nonmicrobial N.
3
2P > 0.10. 4OM apparently digested in the rumen.

*P < 0.05. 5OM truly digested in the rumen.

Journal of Dairy Science Vol. 80, No. 10, 1997


NUTRIENT FLOWS FOR COWS FED GRASS DIETS 2471

TABLE 6. Flow of AA at the duodenum for cows fed unsup- for cows fed the supplemented diet agree with the
plemented grass diet ( U ) or grass diet supplemented with
molassed beet pulp pellets ( S ) . findings of Stakelum et al. ( 2 5 ) who reported in-
creased concentrations of VFA when grass diets were
Treatment
supplemented with barley or beet pulp (molassed or
AA U S P SED1 unmolassed). The molar percentages of VFA did not
(g/d) differ greatly between the treatments. The percentage
Cys 25.8 28.6 NS2 2.01 of acetic acid was somewhat lower, and propionic acid
Asp 219.5 232.5 NS 13.1 somewhat higher, than that previously reported (3, 4,
Met 40.5 40.5 NS 3.5 5 ) for cows fed diets based on grass, which might
Thr 106.1 113.3 NS 6.29
Ser 87.7 94.8 NS 5.75 reflect the effect of season (i.e., the percentage of
Glu 247.1 265.5 NS 14.29 acetic acid is increased, and the percentage of propi-
Gly 145.9 161.7 NS 11.43 onic acid is decreased, as season progresses) (3, 4).
Ala 139.1 150.2 NS 8.16
Val 123.8 133.3 NS 6.54 Sampling was conducted in early and mid May in this
Ile 107.3 115.9 NS 5.35 experiment, which would generally be earlier than
Leu 165.5 176.6 NS 9.29 sampling in other reported work. The most noticeable
Tyr 96.3 102.7 NS 5.75
Phe 107.7 112.2 NS 5.74 difference between the diets in VFA pattern was the
His 43.6 48.2 NS 3.08 rise in the percentage of butyric acid caused by sup-
Lys 148.2 158.9 NS 7.57 plementation. This increase was a feature of high
Arg 88.1 96.6 NS 5.14
Pro 76.8 73.4 NS 6.42 molasses diets ( 2 0 ) and was probably due to the
Total AA 1969 2105 NS 109.3 molasses in the beet pulp in this case, although in a
Total AA N 265 285 previous report, no increase was found in the percent-
AA N, % of total NAN 74.2 73.3
age of butyric acid when a grass diet was sup-
1Standard error of the difference. plemented with molassed beet pulp, but an increase
2P > 0.10. was found with both unmolassed beet pulp and barley
(25).

rumen NH3 concentration decreased when the Rumen OMD


amount of supplement was increased from 1 to 7 kg/d.
The extent of OMD in the rumen was high and was
Reductions that were caused by supplementation not affected by treatment. On average, OMAD was
were probably due to greater availability of energy for
microbial growth and, thus, greater capture and utili-
zation of the available NH3. No study under these TABLE 7. Profile of duodenal AA for cows fed unsupplemented
conditions has measured microbial N synthesis, and, grass diet ( U ) or grass diet supplemented with molassed beet pulp
pellets ( S ) .
in this experiment, the treatments had no significant
effect on microbial N flow. However, microbial N flow Treatment
tended to be higher for cows fed the supplemented AA U S P SED1
diet, which would be consistent with a better capture
( g of AA/100 g of AA)
and utilization of available N.
Cys 1.31 1.35 NS2 0.036
Lactate concentrations were lower than concentra- Asp 11.15 11.05 NS 0.144
tions previously measured in cows fed grass diets (13, Met 2.06 1.92 NS 0.110
25). The lactate concentrations of cows in one of those Thr 5.38 5.38 NS 0.078
Ser 4.45 4.50 NS 0.073
experiments ( 1 3 ) were very high (approximately four Glu 12.56 12.61 NS 0.122
times higher than those measured in this experi- Gly 7.41 7.67 NS 0.307
ment). This difference could be attributed to the Ala 7.06 7.14 NS 0.085
Val 6.29 6.34 NS 0.037
grass in that experiment, which was cut with a flail Ile 5.45 5.51 NS 0.055
type forage harvester. This harvesting method would Leu 8.41 8.39 NS 0.069
have lacerated the grass, which could have caused a Tyr 4.89 4.88 NS 0.042
Phe 5.47 5.33 * 0.040
faster fermentation than that observed in this experi- His 2.21 2.29 NS 0.076
ment. Lys 7.53 7.56 NS 0.070
Total VFA concentrations in the rumen were some- Arg 4.48 4.60 NS 0.169
Pro 3.90 3.51 NS 0.303
what lower in this experiment than in some previous
experiments for cows fed diets based on grass (25, 30, 1Standard error of the difference.
31), but were similar to concentrations reported by 2P > 0.10.
other researchers (3, 4, 5). The higher concentrations *P < 0.05.

Journal of Dairy Science Vol. 80, No. 10, 1997


2472 O’MARA ET AL.

51.4% of OM intake, and OMTD was 74.9% of OM rumen. Values in the range from 68 to 74% have since
intake. When OMD was taken into account, OMAD been reported for perennial ryegrass diets, again at
was 62.9 and 62.8% of digestible OM intake ( DOMI) lower OM intakes than those in the current experi-
for the unsupplemented and supplemented diets, ment (7, 28). The OMTD was 90.5 and 92.7% of
respectively. This result was similar to the conclusion DOMI for the unsupplemented and supplemented
of other researchers ( 2 7 ) based mainly on work with diets, respectively. Thus, very little potentially digest-
steers that had lower OM intakes (27). In that study ible OM escaped digestion in the rumen. These high
(27), 60% of DOMI was apparently digested in the values are supported by results of other researchers,
who have reported even higher values for young
steers fed forage diets [95% ( 3 ) and 97% (4)].

Nutrient Flows
Flows of NAN were higher than N intakes for cows
fed either diet, indicating some recycling of N. There
have been several previous reports (4, 5, 7, 28) of
preduodenal loss of N for cows fed diets based on
grass, but, in those cases, herbage N content and
rumen NH3 concentration (when it was measured)
were higher than those in the current experiment.
The differences in NAN flow between diets were not
significant in this experiment, although flows tended
to be higher, but not significantly higher, for cows fed
the supplemented diet because of increased flow of
microbial protein.
The efficiency of microbial protein synthesis (37.7
and 42.0 g/kg of OMAD for the unsupplemented and
supplemented diets, respectively) was quite high. A
previous review of this subject ( 1 ) concluded that a
value of 32 g/kg of OMAD should be used, although
higher efficiencies for cows fed high quality grass
diets were acknowledged. Microbial N yields of 53.1,
42.4, and 45.3 g/kg of OMAD for cattle fed early, mid,
and late season cuts of perennial ryegrasses, respec-
tively, have been calculated ( 6 ) from previously
reported data (10). Another report ( 3 ) cited mean
values of 56.9, 50.8, and 46.9 g/kg of OMAD for cattle
fed early, mid, and late season cuts of perennial
ryegrasses, respectively. These concentrations are
higher than those in the current experiment, but,
when values are expressed in relation to OMTD, the
differences are smaller. The values from the first
study ( 1 0 ) were 34.7, 29.8, and 31.2 g/kg of OMTD
for early, mid, and late season cuts, respectively, and
the corresponding mean values from the second study
( 3 ) were 35.2, 33.6, and 31.8 g/kg of OMTD for early,
mid, and late season cuts, respectively. The values in
this experiment were 26 and 28.2 g/kg of OMTD for
the unsupplemented and supplemented diets, respec-
tively. The difference between the diets was not
Figure 2. Rumen pH, NH3, and total lactate concentrations at statistically significant but did suggest that the sup-
each sampling time for cows fed the unsupplemented diet ( ♦) and
the diet supplemented with molassed beet pulp pellets ( ⁄) . SED = plement might increase the efficiency of microbial
Standard error of the difference. protein synthesis and N capture in the rumen.

Journal of Dairy Science Vol. 80, No. 10, 1997


NUTRIENT FLOWS FOR COWS FED GRASS DIETS 2473

The efficiency of microbial protein synthesis meas- plementation of high quality grass with molassed beet
ured in this experiment was substantially higher pulp had little effect on the digestion processes and on
than values measured previously with lactating cows the nutrients available for absorption from the small
fed diets of grass silage and concentrates (22), which intestine. The lack of a substantial increase in flow of
ranged from 13.7 to 15.4 g/kg of OMTD. These values nutrients to the duodenum was due to the high sub-
might have been affected by the low DMI in that stitution rate of supplement for forage. There were
experiment, but they may also help to explain the indications that the supplement allowed more effi-
poorer performance of cows fed diets of grass silage cient capture of N in the rumen, which would be
supplemented with moderate amounts of concentrates beneficial for cows fed herbage diets containing high
compared with the performance of cows fed diets N contents (e.g., primary perennial ryegrass growth,
based on grass as noted previously. A large proportion late season perennial ryegrasses, and white clover
of the silage energy that is prefermented to lactate is swards) for which rumen NH3 concentration is very
unavailable for use for microbial growth. high and preduodenal losses of N are observed. In
As with NAN flow, flows of most AA were higher these cases, the supplement could improve the effi-
for cows fed the supplemented diet, although none of ciency of N capture in the rumen and decrease the
the differences were statistically significant. Amino excretion of N into the environment. Whether actual
acid N was 74 and 73% of NAN for the unsup- increases in NAN flow and performance are obtained
plemented and supplemented diets, respectively. depends on the interactions of the supplement with
These values were similar to the value of 70%, which herbage intake and the ability of the cow to respond
has been accepted as a typical value over a range of to increased duodenal NAN supply.
dietary situations (16, 23), but this percentage was
lower than the mean value of 85% reported previously ACKNOWLEDGMENTS
for grass diets fed to sheep (18). However, in that
study (18), the values were expressed as grams of AA The authors express their thanks to Irish dairy
per 100 g of CP flow at the duodenum. This expres- farmers who partially funded these studies through
sion gives a higher value than that when the results the Irish Dairy Levy. We also thank J. J. Cahill, N.
were expressed as grams of AA N per 100 g of N flow Byrne, J. O’Dwyer, M. Kearney, and D. O’Callaghan
at the duodenum, because the latter expression does for care of experimental cows and technical assistance
not account for the increase in mass of AA when and R. Mayes for help with marker analysis.
protein is hydrolyzed. If the calculations of MacRea
and Ulyatt ( 1 8 ) are reworked taking this increase in REFERENCES
mass into account, the mean value is 71%. One other 1 Agricultural Research Council. 1984. The Nutrient Require-
study with cows fed grass ( 3 0 ) reported that AA N ments of Ruminant Livestock. Supplement No. 1. Commonw.
was only 61 to 72% of NAN (30). The reasons for the Agric. Bur., Slough, England.
2 Association of Official Analytical Chemists. 1984. Official
low and variable percentages in that experiment were Methods of Analysis. 14th ed. AOAC, Arlington, VA.
not clear. 3 Beever, D. E., M. S. Dhanoa, H. R. Losada, R. T. Evans, S. B.
The similarity between the profiles of AA for both Cammell, and J. France. 1986. The effect of forage species and
stage of harvest on the processes of digestion occurring in the
diets was not surprising given that a high proportion rumen of cattle. Br. J. Nutr. 56:439.
of the NAN for both diets was of microbial origin. The 4 Beever, D. E., H. R. Losada, S. B. Cammell, R. T. Evans, and
M. J. Haines. 1986. Effect of forage species and season on
profiles differed with respect to many AA from a nutrient digestion and supply in grazing cattle Br. J. Nutr. 56:
previous report in which diets were based on grass 209.
(30), but the profiles were similar to those cited in 5 Beever, D. E., H. R. Losada, D. L. Gale, M. C. Spooner, and
M. S. Dhanoa. 1987. The use of monensin or formaldehyde to
another report (18). control the digestion of the nitrogenous constituents of peren-
nial ryegrass ( Lolium perenne cv. Melle) and white clover
CONCLUSIONS ( Trifolium repens cv. Blanca) in the rumen of cattle. Br. J.
Nutr. 57:57.
6 Beever, D. E., and R. C. Siddons. 1986. Digestion and
These results demonstrate a higher efficiency of metabolism in the grazing ruminant. Page 479 in Control of
microbial protein synthesis than is generally reported Digestion and Metabolism in Ruminants. Proc. VIth Int. Symp.
for diets based on grass silage and concentrates fed at Ruminant Physiol., Banff, AB, Canada. L. P. Milligan, W. L.
Grovum, and A. Dobson, ed. Prentice-Hall, Englewood Cliffs,
similar intakes, which might contribute to the better NJ.
performance of cows fed diets based on grass than of 7 Beever, D. E., D. J. Thompson, M. J. Ulyatt, S. B. Cammell, and
cows fed diets of grass silage supplemented with M. C. Spooner. 1985. The digestion of fresh perennial ryegrass
( Lolium perenne L. cv. Melle) and white clover ( Trifolium
moderate amounts of concentrates ( 5 to 7 kg of DM/ repens L. cv. Blanca) by growing cattle fed indoors. Br. J. Nutr.
d). This experiment also demonstrated that the sup- 54:763.

Journal of Dairy Science Vol. 80, No. 10, 1997


2474 O’MARA ET AL.

8 Boxem, T. J. 1976. Bijvoeding van melkvee in de weide. Publ. rolled barley or sugar cane molasses-based supplements. Anim.
no. 6. Proefstation vour de Rundveehouderij, Lelystad, The Feed Sci. Technol. 50:37.
Netherlands. 21 Morgan, D. J., G. Stakelum, and J. Dwyer. 1989. Modified
9 Boxem, T. J. 1979. Door bijvoeding met krachtvoer lagere neutral detergent cellulase digestibility procedure for use with
grasopname. Bedrijfsontwikkeling 10:53. the ‘Fibertec’ system. Ir. J. Agric. Res. 28:91.
10 Cammell, S. B., D. E. Beever, D. J. Thompson, H. R. Austin, 22 O’Mara, F. P. 1993. Effect of dietary manipulations on milk
H. R. Losada, R. T. Evans, M. C. Spooner, and R. A. Terry. protein yield and concentration and digestion and metabolism
1983. Energy and protein digestion, supply and utilization on of dairy cows. Ph.D. Diss., Univ. College Dublin, Dublin,
two contrasting forages fed to growing steers. Anim. Prod. 36: Ireland.
501.(Abstr.) 23 Rohr, K., P. Lebzien, H. Schafft, and E. Schultz. 1986. Predic-
11 Clancy, M. J., and R. K. Wilson. 1966. Development and appli- tion of duodenal flow of non-ammonia nitrogen and amino acid
cation of a new chemical method for predicting the digestibility nitrogen in dairy cows. Livest. Prod. Sci. 14:29.
and intake of herbage samples. Page 445 in Proc. 10th Int. 24 SAS User’s Guide: Statistics, Version 5 Edition. 1985. SAS
Grassl. Congr., Helsinki, Finland. Assoc. Francaise Prod. Four- Inst., Inc., Cary, NC.
ragere, Inst. Natl. Rech. Agron., Versailles, Cedex, France. 25 Stakelum, G., P. Dillon, and J. Murphy. 1988. The effect of
12 Dillon, P., S. Crosse, G. Stakelum, and F. Flynn. 1995. The concentrate type on the rumen fermentation patterns of grass
effect of calving date and stocking rate on the performance of fed cows and dry matter and crude protein degradability of the
spring-calving dairy cows. Grass Forage Sci. 50:286. herbage. Proc. 12th General Mtg. Eur. Grassl. Fed., Dublin,
13 Dillon, P., G. Stakelum, and J. J. Murphy. 1989. The effect of Ireland. Irish Grassl. Assoc., Belclare, Tuam, Co. Galway,
level of herbage intake and concentrate type on rumen fermen- Ireland
tation pattern, in situ herbage degradability and blood metabo- 26 Taparia, A. L., and A.W.F. Davey. 1970. The effect on food
lite levels in lactating dairy cows. Page 1155 in Proc. 16th Int. intake and milk production of adding concentrates to the ration
Grassl. Congr., Nice, France. Finnish Grassl. Assoc., Helsinki, of pasture-fed cows. N.Z. J. Agric. Res. 13:616.
Finland. 27 Thompson, D. J., and D. E. Beever. 1980. The effect of conserva-
14 Faichney, G. A. 1975. The use of markers to partition digestion tion and processing on the digestion of forages by ruminants.
within the gastrointestinal tract of ruminants. Page 227 in
Page 291 in Digestive Physiology and Metabolism in
Digestion and Metabolism in the Ruminant. I. W. McDonald
Ruminants. Y. Ruckebusch and P. Thivend, ed. MTP Press Ltd.,
and A.C.I. Warner, ed. Univ. New England Publ. Unit., Armi-
Lancaster, England.
dale, New South Wales, Australia.
28 Ulyatt, M. J., D. J. Thompson, D. E. Beever, R. T. Evans, and
15 Hijink, J.W.F. 1978. Supplementary feeding of maize silage to
dairy cows during the grazing season. Publ. no. 12. Proefstation M. J. Haines. 1988. The digestion of perennial ryegrass ( Lolium
voor de Rundveehouderij, Lelystad, The Netherlands. perenne cv. Melle) and white clover ( Trifolium repens cv.
16 Hvelplund, T. 1987. Amino acid content of feed and microbial Blanca) by grazing cattle. Br. J. Nutr. 60:137.
protein and their intestinal digestibility. Page 159 in Proc. CEC 29 Van Soest, P. J., and R. H. Wine. 1967. Use of detergents in the
Sem. Feed Evaluation and Protein Requirement Systems of analysis of fibrous feeds. IV. Determination of plant cell-wall
Ruminants. R. Jarriage and G. Alderman, ed. Comm. Eur. constituents. J. AOAC 50:50.
Commun. Office Official Publ. Eur. Commun., Luxembourg, 30 Van Vuuren, A. M., F. Krol-Kramer, R. A. Van der Lee, and
Luxembourg. H. Corbijn. 1992. Protein digestion and intestinal amino acids
17 Keady, T.W.J., J. J. Murphy, and D. Harrington. 1995. The in dairy cows fed fresh Lolium perenne with different nitrogen
effects of ensiling on dry matter intake and milk production by contents. J. Dairy Sci. 75:2215.
lactating dairy cattle given forage as the sole feed. Grass Forage 31 Van Vuuren, A. M., C. J. Van der Koelen, and J. Vroons-de
Sci. 51:131. Bruin. 1986. Influence of level and composition of concentrate
18 MacRae, J. C., and M. J. Ulyatt. 1974. Quantitative digestion of supplements on rumen fermentation patterns of grazing dairy
fresh herbage by sheep. II. The sites of digestion of some cows. Neth. J. Agric. Sci. 34:457.
nitrogenous constituents. J. Agric. Sci. (Camb.) 82:309. 32 Waghorn, G. C., I. D. Shelton, and V. J. Thomas. 1989. Particle
19 Mason, V. C., S. Beck-Andersen, and M. Rudemo. 1980. breakdown and rumen digestion of fresh ryegrass ( Lolium
Hydrolysate preparation for amino acid determination in feed perenne, L.) and lucerne ( Medicago sativa, L.) fed to cows
constituents. 8. Studies of oxidation conditions for streamlined during a restricted feeding period. Br. J. Nutr. 61:409.
procedures. Z. Tierphysiol. Tierernaehr. Futtermittelkd. 43:146. 33 Zinn, R. A., and F. N. Owens. 1986. A rapid procedure for
20 Moloney, A. P., A. A. Almiladi, M. J. Drennan, and P. J. Caffrey. purine measurement and its use for estimating net ruminal
1993. Rumen and blood variable in steers fed grass silage and protein synthesis. Can. J. Anim. Sci. 66:157.

Journal of Dairy Science Vol. 80, No. 10, 1997

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