Influence of Forage Quality on Rumination Time in Sheep J. G. Welch and A. M. Smith J Anim Sci 1969. 28:813-818.

The online version of this article, along with updated information and services, is located on the World Wide Web at: http://jas.fass.org

www.asas.org

Downloaded from jas.fass.org by on July 17, 2008. Copyright 1969 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.

INFLUENCE

OF

FORAGE QUALITY ON TIME IN SHEEP l

RUMINATION

J. G. WELCH AND A. M. SMITH

University oJ Vermont, Burlington

the major factors determining forONE ofqlTalityfibrous relativein consumption age is its potential. The load the rumen affects the consumption of a given forage; rumination appears to be important in removing indigestible fiber from the rumen by decreasing particle size (Welch, 1967; Pearce and M o l t , 1964). This relationship may explain in part the observation that as a forage matures, the amount that ruminants consume decreases (Blaser, 1964; Reid et al., 1959). Troelsen and Bigsby (1964) showed an association between consumption rate of forages by sheep and time required for grinding to a prescribed fitness in a specially designed recirculating grinder. If forage quality decline is associated with an increased need for remastication to reduce the particle size, the rumination time per unit of forage ingested should increase with declining quality or increasing fiber content. The objective of these experiments was to determine the effect of ingesting single meals of forages of widely different maturities and chemical compositions upon time spent in subsequent rumination. Experimental Procedure Rumination time obtained from a total of 48 individual 24-hr. jaw motion recordings was measured in three experiments. Two groups of four Columbia-Southdale rams weighing between 66 and 94 kg. were used. They were housed in stanchion-type metabolism units and were offered water twice daily. The rams were fed the same medium quality mixed hay during the 2 weeks preceding each experiment and were rested and fed the same mixed hay for at least 7 days between the test feedings. In each experiment the rams were subjected to a 48-hr. roughage withholding period, followed by a single test meal of the forage being studied. The rams were not fed again until 48 hr. after the test meal. Dry matter consumption was equalized
1Unlversity of Vermont Agricultural Experiment Station Journal Article 214. 813

for each forage by offering an initial amount and allowing the rams to eat until the eating pattern subsided on the jaw motion recording. The remaining material was removed and replaced by the same amount of fresh forage (usually less than 100 gm.). The second offering was eaten completely. The particular amounts chosen represented the approximate quantity of hay consumed by the rams during one meal when they were fed on a two-times-a-day schedule. The apparatus attached to the jaw is shown in figure 1. The sensing device was a thin walled rubber tube placed under the jaw of the sheep in such a manner that jaw motion caused compression of the tube and created a pneumatic impulse which was carried through a light plastic tube to the diaphragm of the tambour. The principle is that used by Balch (1952) and also by Oltjen, Sirny and Tillman (1962), but there are important differences. The balloon placed nnder the jaw used by Balch required a positive pressure in the closed system; this causes difficulty when temperature changes produce either expansion or contraction of the air mass in the system immobilizing the recording diaphragm. A very fine leak in the system used in these studies permitted pressure equilibration. The expansible corrugated pneumograph tubing used by Oltjen et al. gave strong accurate impulses, but the tubing was subject to puncture and tearing during long recording periods. In experiment 1 each ram received a single test meal of each of the three chopped forages. Rumination was recorded during the fast, during the single test meal and subsequent rumination period lasting 48 hr., and during the refeeding period. The forages used were a veryearly-cut (prebud) orchardgrass hay, a latecut (full-bloom) mixed grass hay, and oat straw with a high weed content. Test meal consumption of the hays were 792, 790 and 791 gin. dry matter, respectively. The chemical composition of the forages appears in table 1. In experiment 2 a single reversal design

Downloaded from jas.fass.org by on July 17, 2008. Copyright 1969 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.

814

W E L C H AND S M I T H

Figure 1. Halter with attached jaw motion sensing apparatus. Jaw motion caused depression of the thin walled rubber tube, creating a pneumatic impulse which was carried to the recording device by the light plastic tubing. was used with feeding schedules and fast periods similar to those in experiment 1. Two forages, a second cutting mixed alfalfa-grass hay and a relatively weed-free oat straw, were fed as the single test meals, with rumination times recorded as in experiment 1. In experiment 3, rumination of the 1,000 gm. test meals was studied, using the same design and forage materials as in experiment 2, but with 40 gm of cracked corn per head fed daily throughout the entire experiment including the forage fast and test meal periods. Forage protein, crude fiber, ether extract and ash analyses were made according to A.O.A.C. (1960) methods; cell wall constituents (CWC) were determined according to the Van Soest and Wine (1967) method. Analysis of variance was made according to the methods of Steel and Torrie (1960). Figures were plotted using mean rumination time values for 6-hr. periods with period 1 starting at noon. Results a n d Discussion The results of the chemical analysis of the forages appear in table 1. I t is apparent from these data that a wide range of forage composition was used.

Downloaded from jas.fass.org by on July 17, 2008. Copyright 1969 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.

RUMINATION T I M E IN SHEEP
TABLE 1. COMPOSITION OF T E S T FORAGES AS PERCENT OF DRY MATTER Forage Early-cut orchardgrass Late-cut mixed grass Weedy oat straw 2nd cutting mixed alfalfa grass Oat straw % Protein % 22.8 8.0 7.3 17.8 5.4 Crude fiber % Ash % E.E. % 5.2 2,5 2,3 2.6 2.1

815

CWC % 50.7 64.2 67.3 49.5 75.6

Experiment 1 27. I 9,2 36.4 5,7 39.2 6.5 Experiment 2 and 3 28.3 9.1 42.9 9.0

The results of experiment 1 show the general form of the rumination time distributions (figure 2). The first part of the curve shows the normal feeding period (days 1 through 2). The fast started on day 3, resuited in the reduction in rumination activity to almost zero by the first quarter of the fourth day. The test feeding produced a subsequent rapid rise in rumination time, followed by a reduction, again approaching zero as no additional feed was fed. Refeeding during the fourth quarter of day 6 produced a rapid return to normal rumination levels. The summary data in table 2 are the total

of the mean rumination time, following the test feeding and before the refeeding. Figure 3 shows the test feed data shown in figure 2 with an expanded scale used for the X axis. The straw and the mixed grass hay produced the greatest amount of rumination with the peak of rumination activity following the straw feeding occurring later than that for the late cut hay. The early-cut orchardgrass hay produced the least ( p < . 0 5 ) rumination time (table 2). Duration of rumination resulting from the single test meals was similar for all three hays, reaching zero within 48 hr. following consumption of

M A T U R E G R A S S HAY

,K I g T R A w 1. iI

.4

!i l'
li
I , I

II ! I

iI

,,.,.:
V ; "* "

:
f:

ii
,

il

I~l l ,

v i
:

i, .A.I
;r I ..I
;

'

ii
i : :

~ .
I

I!

~o nr -r

.~
I/

,~i ?,
,

I ii EARLY OUT

gl

FAT S,EII~'.,. \!

V \FED ii

XTEST iq
I I I

..~

GRASS HAY

IREFED '~, ~i l , I

DAYS

F i g u r e 2. R u m i n a t i o n time ( h o u r ) in e x p e r i m e n t 1 including n o r m a l r u m i n a t i o n (days 1 a n d 2), f a s t i n g (days 3 a n d 4), r u m i n a t i o n following single test feeding (days 5 a n d 6) a n d r u m i n a t i o n following refeeding (days 7). D a y s are divided into 6-hr. periods, w i t h the first period of each day r e p r e s e n t i n g time b e t w e e n 12 n o o n a n d 6 p.m. E a c h p o i n t indicates average value of four sheep for 6-hr. period.
Downloaded from jas.fass.org by on July 17, 2008. Copyright 1969 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.

816
TABLE

WI,:IA~'H AN1) SM ITH

2. R U M I N A T I O N T I M E S R E S U L T I N G F R O M F E E D I N G S I N G L E MEALS OF FORAGES OF D I F F E R E N T Q U A L I T Y TO FASTED RAMS Test meal weight n Total rumination time Minutes 312 aa 443 aen 513 eb 418 ~ 732 ~ 483 a 793 ~ Rumination ratio Minutes/gm.DM .39 aa .56 a~b .65 eb .45 a .79 e .52 a .85 ~

Forage Early-cut orchardgrass Late-cut mixed grass Weedy oat straw 2nd cutting mixed alfalfa grass Oat straw 2nd cutting mixed alfalfa grass Oat straw 4 4 4 4 4 4 4

as fed gin. 850 825 835 1000 1000 1000 1000

DM gm. Experiment 1 792 790 791 Experiment 2 927 927 Experiment 3 927 927

ab Values within trials with different superscripts are different (p<.05). ae Values within trials with different superscripts are different (p<.O1).

the test meal. In experiment 2, the straw produced the greater ( p < . 0 1 ) rumination time when compared to the second cutting mixed hay. Again the peak resulting from the straw feeding was later than the peak resulting from the hay feeding (figure 4). Similar results were observed in experiment 3 (table 1 and figure 5) in which grain was fed. The second cutting mixed hay produced the lowest (p<.01) total rumination time. Rumination in the straw-fed animals again

reached a peak 18 hr. after the test feeding. Association between the fiber fractions of the hays fed and the total rumination time produced by the test feedings was high. The simple correlation between mean rumination time and CWC intake in the three trials was 0.99 (figure 6). One feeding of corn (500 gm). was included in the CWC and crude fiber intake calculations in experiment 3, using the value of 10.4% CWC for cornmeal reported by Van Soest and Wine (1967),

il MATURE GRASS HAY

3
I
I

!.,.1 ~
I /,'
/

,,',,
~ ~,

STRAW

i
2

"~ I
"~l

I'

iI

'~

,,"

~EARLY

OUT GRASS HAY

g,,

II

PERIOD
F i g u r e 3. R u m i n a t i o n t i m e ( h o u r s ) f o r t e s t f e e d i n g s in e x p e r i m e n t 1. E a c h p e r i o d r e p r e s e n t s 6-hr. w i t h p e r i o d 1 f o r e a c h d a y s t a r t i n g at 12 n o o n . E a c h p o i n t i n d i c a t e s a v e r a g e v a l u e of f o u r s h e e p f o r 6-hr. period.

Downloaded from jas.fass.org by on July 17, 2008. Copyright 1969 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.

R U M I N A T I O N T I M E IN SHEEP A t \

817

2ND CUT HAY

/

//

\ / \
\

STRAW

A
,

jl

ii

\\

\\

%1.
o T

/;' l /
I I I

X",,
I I i ~, I v

/
I I

4
PERIOD

Figure 4. Rumination time (hours) for test feeding in experiment 2. Each period represents 6-hr. with period 1 for each day starting at 12 noon. Each point indicates aveiage value of four sheep for 6-hr. period.

and the value of 2.3% for crude fiber (N.R.C., 1966). The simple correlation coefficient between mean rumination time and crude fiber intake was also 0.99. Pearce (1965) suggested that increased protein intake could decrease rumination time of sheep.

In these experiments, although the protein effect could be a factor, the simple correlation between protein intake and total rumination time was only 0.24. Weston (1968) found that straw was retained in the rumen longer than better quality forage material.

3.

/^\ ~,.,.STRAW 2NDCUTHAY / "~

//,
~
I

2.

I 9

/,.-~
0

',

D
I I I I I

4 1 PERIOD

Figure 5. Rumination time (hours) resulting from the test feeding in experiment 3. Each period represents 6-hr. with period 1 for each day starting at 12 noon. Each point indicates average value of four sheep for 6-hr. period.
Downloaded from jas.fass.org by on July 17, 2008. Copyright 1969 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.

818
800

W E L C H AND S M I T H
r=.99

6OO

uJ

u~400 z X2o (
o

100 200 3 0 0 4 0 0 5 0 0 6 0 0 7 0 0 8 0 0 OWC INTAKE g

Figure 6. Relation of minutes of rumination time to cell wall constituent intake in sheep fed single m e a l s of r o u g h a g e s of different quality. G r o u n d s t r a w w a s c o n s u m e d a t h i g h e r levels a n d p r o d u c e d less r u m i n a t i o n t h a n t h e Association of Official Agricultural Chemists. Washington, D. C. Balch, C. C. 1952. Fatcors affecting the utilization of food by dairy cows. 6. The rate of contraction of the reticulum. British J. Nutr. 6:366. Blaser, R. E. 1964. Symposium on Forage Utilization: Effects of fertility levels and stage of maturity on nutritive value. J. Animal Sci. 23:246. N.R.C. 1966. Nutrient requirements of domestic animals, No. 3. Nutrient requirements of dairy cattle. National Research Council, Washington, D. C. Oltjen, R. R., R. J. Sirny and A. D. Tillman. 1962. Purified Diet Studies with sheep. J. Animal Sci. 21: 277. Pearce, G. R. and R. J. Moir. 1964. Rumination in sheep. I. The influence of rumination and grinding upon the passage and digestion of food. Australian J. Agr. Res. 15:635. Pearce, G. R. 1965. Rumination in sheep. II. The circadian pattern of rumination. Australian J. Agr. Res. 16:635. Reid, J. T., W. K. Kennedy, K. L. Turk, S. T. Slack, G. W. Trimberger and R. P. Murphy. 1959. Effect of growth stage, chemical composition and physical properties upon the nutritive value of forages. J. Dairy Sci. 42:567. Steel, R. G. D. and J. H. Torrie. 1960. Principles and procedures of statistics. McGraw-Hill Book Company, Inc. New York, N. Y. Troelsen, J. E. and F. W. Bigsby. 1964. Artificial mastication--a new approach for predicting voluntary forage consumption by ruminants. J. Animal Sci.

chopped form. (Weston and Hogan, 1967). Preventing rumination by muzzling caused a reduced rate of passage of forage material
from the rumen (Pearce and Moir, 1964).

Feeding ground roughage to the muzzled sheep allowed for normal rates of passage, These results indicate that rumination is an important process in promoting passage of
coarse fibrous materials from the rumen, and may be an important f a c t o r in l i m i t i n g in-

take of poor quality forages. Fibrous m a s s appears to be the important variable or closely associated with the important variable
in c o n t r o l l i n g r u m i n a t i o n t i m e u n d e r t h e single m e a l f e e d i n g c o n d i t i o n s of t h e experim e n t s r e p o r t e d here.

Summary The effect of different forages on the rumination time in two groups of four rams was studied in 3 experiments. The test forages were fed to fasted rams in single equal dry matter meals and the rumination time recorded. Poor-quality roughage with high fibre and cell wall constituent c o n t e n t p r o duced the greatest amount of rumination, Combining the mean rumination times for all 3 experiments, 1.05 min. rumination time w a s required for each gram of cell wall constituents ( r ~ 0 . 9 9 ) and 1.63 rain. of rumination time was required for each gram of crude
fiber i n g e s t e d ( r z 0 . 9 9 ) .

23:1139.
Van Soest, P. J. and R. H. Wine. 1967. Use of detergents in the analysis of fibrous feeds. IV. Determination of plant cell-wall constituents. ~'. Assn. Official Agr. Chem. 50:50. Welch, J. G. 1967. Appetite control in sheep by indigestible fibers. J. Animal Sci. 26:849. Weston, R. H. and ~'. P. Hogan. 1967. The digestion of chopped and ground roughages by sheep. I. The movement of digesta through the stomach. Australian J. Agr. Res. 18: 789. Weston, R. H. 1968. Factors limiting the intake of feed by sheep. III. The mean retention time of feed particles in sectons of the almentary tract. Australian J. Agr. Res. 19:261.

L i t e r a t u r e Cited
A.O.A.C. 1960. Official Methods of Analysis (9th ed.).

Downloaded from jas.fass.org by on July 17, 2008. Copyright 1969 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.