Agropecuaria-UNNE

*6.03-R02234* S - 6 3 6 . 0 8 4 M C C ej.1

Methane production by steers on pasture

W. R McCaughey , K. Wittenberg , and D. Corrigan
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^Agriculture and Agri-Food Canada, Research Centre, P.O. Box 1000 A, Brandon, Manitoba, Canada R7A 5Y3; ^Department of Animal Science, University of Manitoba, Winnipeg, Manitoba, Canada R3T 2N2. Received 2 December 1996, accepted 21 July 1997.

McCaughey, W. P., Wittenberg, K. and Corrigan, D. 1997. Methane production by steers on pasture. Can. J. Anim. Sci. 77: 519-524. In order to determine the quantity of methane (CH ) produced by steers on pasture, 16 steers with a mean weight of 356 25 kg were randomly selected from a larger group of cattle (n = 48) to evaluate the effects of grazing management and monensin controlled release capsule (CRC) administration on ruminai C H production using the sulphur hexafluoride (SF ) tracer-gas technique. Pasture management treatments consisted of two grazing systems (continuous stocking or 10-paddock rotational stocking) at each of two stocking rates (low, 1.1 steer h a or high, 2.2 steers h a ) with two replications of each pasture treatment. Half of the animals on each pasture treatment were administered a monensin CRC delivering 270 mg d , and untreated animals served as controls. During, the 140-d grazing season, one steer from each treatment-replicate combination was sampled to determine daily intake and C H production on four occasions. The chemical composition of diets differed between grazing management treatments and sampling periods. Voluntary intake and C H production, adjusted for differences in body weighty were unaffected by grazing management, sampling period or by monensin CRC administration and averaged 0.69 0.1 L kg B W d across all grazing management treatments. The energy lost through eructation of C H averaged 4.5 1.4% of gross energy intake.
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Key words: Methane, cattle, environment, digestion efficiency, pasture, forage McCaughey, W. P., Wittenberg, K. et Corrigan, D. 1997. Production de mthane par les bouvillons au pturage. Can. J. anim. Sci. 77: 519524. Afin de dterminer la quantit de mthane ( C H ^ dgage par les bouvillons l'herbe, nous avons pris au hasard parmi un groupe de 48 16 bouvillons d'un poids moyen de 356 25 kg. L'objet tait d'valuer au moyen d'un gaz marqueur, l'hexaflorure de soufre (SF ) les effets de la conduite du pturage et de l'administration d'un diffuseur de monensin (DM) sur la production ruminale de C H . Les traitements de conduite de la pture, deux rceptions, comportaient deux systmes : pturage continu ou pturage tournant 10 parcs, chacun 2 taux de chargement, l'un bas, 1,1 bouvillon l'autre 2,2 bouvillons par hectare. La moiti des animaux affects chaque traitement recevaient le diffuseur de monensin, rgl de faon librer 270 mg j, l'autre moiti servant de tmoin. Durant les 140 j de la saison de pture, un bouvillon de chaque combinaison traitement-rptition tait choisi pour mesurer quatre reprises la prise alimentaire et la production de mthane. La composition chimique du fourrage diffrait selon les modes de conduite du pturage et les priodes de prlvement. L'ingr libre et la production de C H , corrigs en fonction des diffrences de poids corporel, n'taient modifis ni par le mode de conduite, ni par la priode de prlvement, ni par la prsence du monensin. La production de mthane tait en moyenne de 0,69 0 , 1 L kg PC" j , tous modes de conduite du pturage confondus. Les penes d'nergie par ructation de C H se situaient autour de 4,5 1,4 % de l'ingr nergtique brut.
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Mots cls: Mthane, bovin, environnement, efficacit digestive, pturage, fourrage

The dietary, environmental and microbiological aspects of methane (CH ) production in ruminants have been recently reviewed (Johnson and Johnson 1995; McAllister et al. 1996). The production of C H by cattle and other ruminants has become a subject of scientific debate as awareness and concern over global warming increase. Domestic animals (primarily cattle) may be responsible for as much as 15% of the world's output of C H (Crutzen et al. 1986) and, therefore, may be considered as significant contributors to the greenhouse effect. Other significant contributors of C H are natural wetlands (21%), rice fields (20%), losses from fossil fuel production (14%), burning of biomass (10%) and landfills (7%). At the present time, CO-, is considered to be the most important greenhouse gas. However, atmospheric C H concentrations are increasing rapidly. This is causing concern since C H is approximately 30 times more efficient than C 0 in trapping heat. In time, C H may become the dominant greenhouse gas (Cicerone and Oremland 1988; Pearce 1989).
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Methane production by ruminant animals results from the fermentation of feeds tuffs in the gastrointestinal tract and represents a loss of dietary energy. Most studies have measured C H gas production using respiration chambers in which animals are fed conserved, and often processed, forages (Blaxter and Clapperton 1965). The majority of work in this field has been conducted on dairy cattle fed primarily high energy rations (Holter and Young 1992). In contrast to dairy cattle, most beef cattle are kept on pasture for 5 to 12 mo each year, depending on the geographic location. Differences between confined, chamber-fed animals consuming conserved feedstuffs and grazing animals in terms
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A b b r e v i a t i o n s : ADF, acid detergent fibre; ADG, average daily gain; C H , methane, CP, crude protein; CRC, controlled release capsule; DMI, dry matter intake; GE, gross energy; GEI, gross energy intake; IVOMD, in vitro organic matter digestibility; NDF, neutral detergent fibre; OMI, organic matter intake, SF , sulphur hexafluoride
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of feeding behaviour and rumen digestion kinetics may be expected as animals on pasture may have higher voluntary intake and rate of passage than chamber-fed animals. Pastured animals may also graze for extended periods which may total as long as 812 h per day (Popp 1995). Ionophores, such as the commercial feed additive monensin have been reported to reduce C H production during digestion (Schelling 1984). The recent development of a controlled release capsule (CRC; Provel/Elanco, Guelph, ON) to meter out monensin intra-ruminally makes it theoretically possible to reduce C H production in free-ranging cattle without the need for supplemental feeding. Johnson et al. (1994) described a new technique for measurement of C H production by cattle using an inert gas tracer source placed in the rumen of the animal. Until this S F tracer gas technique was developed it was impossible to obtain good estimates of C H production by pastured ruminants.
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Currently, no published information exists documenting C H production under range or pasture conditions. The objectives of the current experiment were to measure C H production by steers grazing alfalfa/grass pastures and to observe the effects of grazing management strategies (grazing system and stocking rate) and ionophore use on C H production.
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MATERIALS AND METHODS During the 1994 grazing season, 16 crossbred yearling (13 mo) steers averaging 356 25 kg were randomly selected from a larger group of 48 similar animals for determination of C H production during the grazing season. The experiment was conducted on eight, 3.7-ha pastures which had been used in a grazing system and stocking rate trial for the previous 3 yr. All eight pastures were of similar botanical composition; based on basal area measurements, pasture botanical composition was 60.0 5.3% alfalfa (Medicago
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way through each of the two rotations and would give the best approximation of average forage quality in rotationally grazed pastures. This created a structure which allowed the effects of grazing system, stocking rate and monensin CRC bolus to be determined. The experimental protocol was reviewed by the Brandon Research Centre Animal Care Committee to ensure that animals used in this study were cared for in accordance with the principles outlined by the Canadian Council on Animal Care. Four esophageally fistulated steers were kept on pastures of similar botanical composition to the treatment pastures. On each sampling date, one esophageally fistulated steer fitted with a canvas collection bag was used to obtain diet quality samples for each pasture. The fistulated steers were fasted overnight prior to being used to sample each pasture treatment. Four of the pastures were sampled in the morning and four were sampled in the afternoon. The fistulated steers were kept in the treatment pastures for 20 to 40 min to collect sufficient plant material for chemical analysis following the protocol of Cohen (1979). Extrusa samples were squeezed through two layers of cheesecloth and saliva fractions were collected and used to correct analyses for organic matter losses (Cohen 1979). Squeezed samples were dried in a forced draught oven at 50C for at least 48 h and ground using a Wiley mill fitted with a 1-mm screen. Dried samples were analyzed for CP (Association of Official Analytical Chemists [AOAC] 1984, method 7.025), ADF and NDF (Goering and Van Soest 1970) and ash (AOAC 1984, method no. 7.009). GE was determined using a Parr adiabatic bomb calorimeter (model 1241). IVOMD was determined by a modification (Troelsen and Hanel 1966) of the method of Tilley and Terry (1963) using bovine instead of ovine inoculum. Voluntary intake was determined using chromium sesquioxide (Cr-,0 ) controlled release capsules (Captechrome, Nufarm Ltd.. New Zealand) to estimate fecal output (Barlow et al. 1988). The IVOMD values of pasture samples were used to predict voluntary intake of digestible organic matter. To allow for the lag times associated with passage of feed through the digestive tract and to ensure that intake estimates approximately corresponded to the C H collection periods, fecal samples were taken 48 h after paddock entry and 12 h after paddock exit. Methane gas production was determined by the S F tracer gas method (Johnson et al. 1994). Using this method, permeation tubes (1.25 x 4 cm) containing S F with a pre-determined release rate were placed in the rumen. The rate of S F release from a permeation tube was determined by measuring the weight loss of the permeation tube over a set period of time during which the permeation tube was incubated at 39C. Release rates of the permeation tubes used in this study ranged from 356 to 1276 ng m i n . Expired gases were sampled using a gas collection apparatus fitted to a halter and suspended from the neck of the animal (Fig. 1). This apparatus consisted of a 13-cm-diameter stainless steel sphere evacuated to approximately 30 mm Hg. The sphere was suspended by a neck-strap from the neck of the animal and was attached to a halter fitted with an air-tight connection to a 90-cm length of restriction tubing
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sativa L.), 28.6 5.4% meadow bromegrass (Bromus bieber-

steinii Roem and Schult.), and 4.9 0.6% Russian wildrye (Psathyrostachys juncea (Fisch.) Nevski) with the balance being made up of quackgrass {Elytrigia repens (L.) Nevski) and other weeds. The experimental treatments consisted of four combinations of two grazing systems (continuous stocking or 10-paddock rotational stocking) and two stocking rates (low, four steers per pasture stocked at 1.1 steers h a ; or high, eight steers per pasture stocked at 2.2 steers h a ) , with two replications of each pasture treatment. Within each pasture, half of the animals received a monensin CRC and the remainder were untreated. One steer receiving a monensin CRC bolus and one untreated steer were selected at random from each pasture to monitor C H production on four occasions during the 1994 grazing season.
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The first two sampling dates occurred during the first grazing rotation when steers entered (22 June) and exited (27 June) the fifth paddock in a ten-paddock rotation. The third and fourth sampling dates occurred when steers on rotationally grazed pastures entered (3 August) and exited (6 August) paddock five during the second grazing rotation. Continuously grazed pastures were sampled concurrently with rotationally grazed pastures. The fifth paddock was selected for sampling because it was approximately half

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(5/1000" i.d.) with an in-line 15 um filter and flexible nose piece. Gas collection spheres were changed approximately every 8 h. Based on preliminary research, expired gases were sampled in a linear manner over an 8-h period. Prior to gas sampling, the animals were allowed to adapt to the gas collection apparatus. No effect was observed in grazing behaviour once the test animal was adapted to the apparatus. Following the removal of the gas collection spheres from the steers, they were pressurized to 16 psi with pure N, and taken to the laboratory for further analysis. The pressurization was to prevent sample contamination prior to gas analysis and to allow injection of a gas sample into the sample loop of a gas chromatograph. A gas chromatograph (Star 3000, Varian, Mississauga, ON) fitted with electron capture and flame ionization detectors was used for determination of S F and CH , respectively. The samples were run on the gas chromatograph using a Molecular Sieve 5A (180 cm) column and a Poropak QS (180 cm) column for S F and C H , respectively. The temperature of the column oven was 35C and nitrogen was used as the carrier gas with a flow rate of 30 mL min . Prepared standards were used to standardize the gas chromatograph for both S F (Scott-Marrin Inc., Riverside, CA) and C H (Supelco, Mississauga, ON) prior to sample analysis. Methane production was determined using the formula developed by Johnson et al. (1994):
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where Y- = methane production of the ijklth steer, 7J = i'th grazing treatment, R- = jth replication, M = kth monensin treatment, P, = /th sampling period and e- = the experimental error term. Grazing treatments were tested against the error A term ((77?)y), the monensin effect and its interaction with main plot effects were tested against the error B term ((7i?M) ), all time effects and their interactions were tested against the experimental error term ( ^ i m ) - When errore and B terms were found not to differ from the experimental error term, all three error terms were pooled for the final analysis.
k[ k k[ ijk

RESULTS
Diet Q u a l i t y

C H production (ng m i n ) = Perm, tube S F release rate (ngmm-VtCH^/tSFJ

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Both grazing management and sampling period had significant effects (P < 0.05) on the nutritive value of forage ingested by steers (Table 1). Interactions between grazing treatment and sampling period were significant for CP, IVOMD, ADF and NDF (Tables 1 and 2). There were no effects of grazing management treatments on CP, IVOMD, ADF or NDF content on the 22 June sampling date (Table 2). In contrast, grazing management had significant effects (P < 0.05) on nutritive value of ingested forage on each of the last three sampling dates. However, the pattern observed was not consistent between sampling periods or dietary parameters. Diet quality was often greater (P < 0.05) during June than it was during August; however, this was dependent somewhat on the grazing treatment used. Neither sampling period nor grazing management appeared to affect ash content or GE value of the ingested forage.
Performance, Intake a n d C H
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The experiment was analyzed using a split-split plot design with the main plots arranged in a completely randomized design. The analysis was performed using the GLM procedure of the SAS Institute, Inc. (1985). The main plot treatments were the four possible combinations of grazing system and stocking rate, sub-plots were the monensin treatments and sub-sub-plots were the sampling dates. The statistical model used was therefore as follows: *m =
r

Production

V ()ii

W i k

(a)**

Pi

W4MTpV(Aki

+ e ijkl

Grazing management and sampling period affected average steer weight and C H production (Table 3). Grazing management also affected ADG. Average steer weights differed among treatments and ranged from 380.1 kg for steers continuously grazed at high stocking rates to 417.1 kg for steers rotationally grazed at low stocking rates (Table 4). Steers on low stocking rate treatments were heavier (P < 0.05) than steers on high stocking rate treatments for both grazing systems. The ADG of steers over the grazing season was great-

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Table 1. Analysis of variance for diet quality factors (DM basis) Parameters CP (%) IVOMD (%) ADF (%) NDF (%) Ash (%) GE (Meal kg" )
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P ***
***

TxP

cv
0.83 0.76 0.83 0.81 0.22 0.31 17.5 8.2 13.8 15.2 51.0 7.4

SD 2.9 4.9 5.1 8.3 6.2 0.3

Mean 16.6 59.3 37.1 54.5 12.2 4.21

*** NS **
** +

NS NS

*** NS NS

* ** *** *** NS NS

T = grazing treatment; P = sampl ng period. P < 0.05, P < 0.01, and P < 0.001, respectively; NS , not significant. Table 2. Effect of grazing system (rotational and continuous stocking), stocking rate (Hi-SR, 2.2 steers h a ; Lo-SR, 1.1 steers h a ') and stage of season on chemical composition (DM basis) of esophageal extrusa samples from yearling steers grazing alfalfa/grass pastures
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Rotational stocking Parameter CP (%) Period 22 Jun. 27 Jun. 3 Aug. 6 Aug. SEM P-value 22 Jun. 27 Jun. 3 Aug. 6 Aug. SEM P-value 22 Jun. 27 Jun. 3 Aug. 6 Aug. SEM f-value 22 Jun. 27 Jun. 3 Aug. 6 Aug. SEM P-value Avg.
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Continuous stocking Hi-SR 25.4a// 24.8aA \4.9aB \33aB 1.3 0.0001 61.laA 69. la A 41.1bB 53.6a5 5.6 0.0077 23.7a5 23.7c5 423aA 4%.\aA 3.0 0.0001 31.0a5 34.065 62.\aA
66.30A

Hi-SR 21.8aA 15.065 l.laf 10.765 2.0 0.0023 6%.7aA 56.5bC 62.5aB 53.5aC 1.3 0.0001 27.6aC 42.2aA 33.865 46.3aA 2.5 0.0001 44.1a5 64.7aAB 50AbAB l\.6aA in 0.0215 12.0 4.22

Lo-SR 22.8aA 19.66/ \1.9aA 9.5bcB 2.6 0.0014 67.0aA 64.1 abA 62 A aA 49.9aB 2.0 0.0001 26.5a5 31.965 30.565 52.1 aA 4.3 0.0003 38.4a5 45.365 42.565 lO.laA 6.8 0.0021 13.1 4.17

Lo-SR 20.2aA \5.2bA 9.065 8.3c5 2.4 0.0009 66.9a^ 60.4a65 53.7a6C 50.8aC 2.6 0.0008 3I.la5 3&.4aAB 4%.2aA
48.0.4

SEM 1.5 1.2 0.7 0.4

P- value NS 0.0040 0.0002 0.0002

IVOMD (%)

1.4 1.5 2.4 1.0

NS 0.0212 0.0275 NS

ADF (%)

2.0 1.4 1.7 2.0

NS 0.0002 0.0007 NS

4.2 0.0037 48.2o5 62.8^ 722aA Ti3aA 6.3 0.0066 10.5 425 4.5 3.5 2.4 1.2 NS 0.0021 0.0002 0.0747

NDF (%)

3.9 0.0001 13.4 4.19

Ash (%) GE (Meal kg" )

2.6 0.11

NS NS

Avg.

ac Values within grazing treatments with different letters differ (P < 0.05) using Student-Newman-Keul's test. ACValues within sampling periods with different letters differ (P < 0.05) using Student-Newman-Keul's test.

est (1.48 kg d ) for steers on lightly stocked, continuouslygrazed pastures and lowest (1.07 kg d ) for steers on heavily stocked, continuously grazed pastures. Steers on rotationally stocked pastures at both high and low stocking rates (1.26 and 1.29 kg d , respectively) were intermediate in terms of ADG. Neither grazing management nor sampling period affected ste_er dry matter intake (x = 13.8 kg), organic matter intake (x = 12.5 kg) or gross energy intake (x = 58.8 Meal d-') (Table 3).
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Methane production was greatest for steers continuously grazed at low stocking rates and least for steers continuously grazed at high stocking rates (306.7 vs. 242.2 24.2 L (H; P < 0.05) Table 4). Methane production by rotationally grazed steers on heavily and lightly stocked pastures was intermediate between these two values and did not differ from either (263.8 and 280.1 24.2 L d~ , respectively). Methane production per ha was calculated as 540.4, 302.8,
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523.6 and 331.6 L h a d for rotationally grazed-heavily stocked, rotationally grazed-lightly stocked, continuously grazed-heavily stocked and continuously grazed-lightly stocked treatments, respectively. However, when C H production measurements were compared on a per kilogram of body weight basis, no differences were found between grazing treatments, and it averaged 0.69 0.06 L kg B W d (Table 3). The average energy lost by steers grazing alfalfa/grass pastures through eructation of C H was not affected by grazing management treatments and was estimated at 4.5 1.4% of gross energy intake. No benefit was observed from administration of monensin CRC in terms of animal performance, voluntary intake or C H production.
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DISCUSSION The S F technique allows direct estimates of ruminai C H production to be made in free ranging cattle, but the tech6 4

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Table 3. Analysis of variance for animal performance, intake and methane production
Parameter T M NS NS NS NS NS NS NS NS NS NS NS NS NS NS NS NS P TxP NS

MxTxP NS

CV 4.0 13.2 19.5 18.9 19.7 21.7 22.6 23.1

SD 15.9 0.02 2.7 2.4 11.6 59.3 0.15 2.0

Mean 397.8 1.27 13.9 12.5 58.8 273.2 0.69 4.5

*** Wt. (kg) ADG (kg d-') *** DM Intake (kg d~') NS OM Intake (kg d~') NS GE Intake (Meal d" ) NS Methane (L d ) * Methane (L kg BW"> d" ) N S Methane (% o f G E I ) NS
1 _ l 1

***

NS

NS NS NS * NS NS

NS NS NS NS NS NS

NS NS NS NS NS NS

NS NS NS NS NS NS

0.90 0.70 0.72 0.72 0.73 0.61 0.48 0.82

T = grazing treatment; M = monensin treatment; P = sampling period. *, *****P < 0.05, P < 0.01, P < 0.001, respectively; NS, not significant.

Table 4. Effect of grazing system (rotational and continuous stocking), stocking rate (Hi-SR, 2.2 steers h a ; Lo-SR, 1.1 steers h a ) on animal performance, intake and methane production by yearling steers grazing alfalfa/grass pastures
Rotational stocking Parameter Weight (kg) ADG (kg d" ) DM Intake (kg t H ) OM Intake (kg d" ) GE Intake (Meal d~') Methane (L d ) Methane (L kg BW~ <H) Methane ( % o f G E I )
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Continuous stocking High 380.1c 1.07c 13.51 12.17 57.36 242.26 0.64 4.4 Low 402.66 1.48a 13.20 12.09 56.16 306.7a 0.77 5.2 SEM 3.9 0.1 0.88 0.91 4.23 24.2 0.06 1.4 P-value 0.0001 0.0001 NS NS NS 0.0325 NS NS

High 391.56c 1.26* 14.94 13.29 62.57 263.8a6 0.67 4.1

Low 417.1a 1.296 13.61 12.40 58.25 280. lab 0.67 4.3

ac Values within the same row that are followed by different letters are significantly different (P < 0.05) using Student-Newman-Keul's test.

nique does not allow quantification of C H produced in the hind-gut and lost via the rectum. Therefore, to estimate total methane produced by the animal, values will need to be adjusted upward to account for the proportion of C H produced in the hind-gut. Unfortunately, there are no good estimates of hind-gut C H production in the literature for cattle and this is an area which requires research attention. The results of this experiment indicate that diet quality of steers grazing alfalfa/grass pasture changed dramatically over a grazing season in continuously grazed pastures, and from paddock entry to paddock exit for rotationally grazed pastures. This resulted in the complex pattern of interactions seen in Table 2. Thus, the experiment was successful in creating a wide range of diet qualities. However, despite the wide range in diet quality, no differences in voluntary intake or C H production per unit weight of steer (L kg B W d ) were observed and C H production could not be predicted on the basis of intake and diet quality. These results are similar to recent observations in respiration chamber experiments (G. Mathison, personal communication) where dry matter intake has been the primary determinant of C H production. Quality factors such as digestibility have been of secondary value for predicting C H production. This is also supported by the work of Blaxter and Clapperton (1965) where both feed intake and digestibility were found to be factors affecting C H production. These respiration chamber experiments of Blaxter and Clapperton (1965) were tightly controlled with animals being fed diets of consistent quality and quantity. In contrast, cattle on pasture can graze selectively and there may be more variability associated with the measurement of intake and diet quality than is the case for chamber-fed animals. As no differences in voluntary intake
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were observed in the current experiment due to changes in either grazing system or stocking rate it is not surprising that C H production by cattle on pasture could not be predicted on the basis of intake and diet quality. Although C H production ranged from 242.1 to 306.7 L d depending on grazing treatment, no differences were observed when these values were corrected for differences in body weight. Overall, the energy lost through eructation of C H averaged approximately 4.5 1.4%. This value is within the range of values reported for forage-fed, non-lactating dairy cows and heifers (Holter and Young 1992) but is slightly lower than the values reported by Johnson and Huyler (1994) for growing beef steers and heifers. These results are not unexpected, given the high levels of intake observed in this experiment. However, no direct measurements of C H production by free ranging cattle have been published previously.
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The intake values reported in this experiment are higher than those commonly observed in grazing experiments using grass-based pastures (Minson and Wilson 1994), but are similar to observations reported previously on the same alfalfa-grass pastures (Popp 1995). Therefore, some care should be used in extrapolating values obtained in this experiment directly to situations where grasses are the predominant forage species. Also, production of C H reported in this experiment is higher than other estimates reported in the literature (Crutzen et al. 1986). This can be explained by the higher voluntary intakes obtained on legume-based compared to grass-based pastures (Minson and Wilson 1994) as digestibility and rate of passage are greater for legumes than grasses. The monensin CRC used in this experiment released monensin at a rate of 270 mg d (Monensin CRC
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technical manual). It is possible that rumen levels may have been inadequate to lower steer C H production. Based on the high voluntary intake levels observed, it is likely that monensin was only present in the rumen at 18 to 21 ppm concentrations. This is below the level commonly recommended in feedlot diets (33 ppm) and consequently may not have been effective at the release rate used.
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The results obtained in this study indicate that it will not be easy to manipulate C H production of steers grazing alfalfa/grass pastures through changes in grazing management. On the improved pastures used in this experiment, C H production and voluntary intake remained relatively constant regardless of variations in diet quality.
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Crutzen, P. J., Aselmann, I. and Seiler, W. 1986. Methane production by domestic animals, wild ruminants, and other herbivorous fauna, and humans. Tellus 38B: 271-284. Goering, H. K. and Van Soest, P. J. 1970. Forage fiber analysis. Agricultural Handbook No. 379. USD A, Washington, DC. Hotter, J. B. and Young, A. J. 1992. Methane prediction in dry and lactating holstein cows. J. Dairy Sci. 75: 2165-2175. Johnson, K. A. and Johnson, D. E. 1995. Methane emissions from cattle. J. Anim. Sci. 73: 2483-2492. Johnson, K., Huyler, M., Westberg, H., Lamb, B. and Zimmerman, P. 1994. Measurement of methane emissions from ruminant livestock using a S F tracer technique. Environ. Sci. Technol. 28: 359-362. McAllister, T. A., Okine, E. K., Mathison, G. W. and Cheng, K. J. 1996. Dietary, environmental and microbiological aspects of methane production in ruminants. Can. J. Anim. Sci. 76: 231-243. Minson, D. J. and Wilson, J. R. 1994. Prediction of intake as an element of forage quality. Pages 5 3 3 - 5 6 3 in G. C. Fahey, ed. Forage quality, evaluation and utilization. American Society of Agronomy, Madison, WI.
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ACKNOWLEDGEMENTS The technical assistance of Clayton Robins, Cindy Rose and Jim McGhee is gratefully acknowledged. This work was funded by Agriculture and Agri-Food Canada's Greenhouse Gas Initiative. Association of Official Analytical Chemists. 1984. Official methods of analysis. 14th ed. AOAC, Washington, DC. Barlow, R., Ellis, K. J., Williamson, P. J., Costigan, P., Stephenson, P. D., Rose, G. and Mears, P. T. 1988. Dry matter intake of Hereford and first cross cows by controlled release of chromic oxide on three pasture systems. J. Agric. Sci. (Camb.) 110:217-231. Blaxter, K. L. and Clapperton, J. L. 1965. Prediction of the amount of methane produced by ruminants. Br. J. Nutr. 19: 511-522. Cicerone, R. J. and Oremland, R. S. 1988. Biogeochemical aspects of atmospheric methane. Global Biogeochem. Cycles 2: 299-327. Cohen, R. D. H. 1979. Factors influencing the estimation of the nutritive value of the diet selected by cattle fistulated at the oesophagus. J. Agric. Sci. (Camb.) 93: 607-611.

Pearce, F. 1989. Methane: the hidden greenhouse gas. New Scientist. May 6: 3 7 - 4 1 . Popp, J. D. 1995. Beef production on alfalfa grass pastures. Ph.D. Thesis. University of Saskatchewan, Saskatoon, SK. 185 pp. SAS Institute, Inc. 1985. SAS user's guide: Statistics. Version 5 ed. SAS Institute, Inc., Cary, NC. 956 pp. Schelling, G. T. 1984. Monensin mode of action in the rumen. J. Anim. Sci. 58: 1518-1527. Tilley, J. M. A. and Terry, R. A. 1963. A two stage technique for the in vitro digestion of forage crops. J. Br. Grassl. Soc. 18: 104-111. Troelsen, J. E. and Hanel, D. J. 1966. Ruminant digestion in vitro as affected by inoculum donor, collection day and fermentation time. Can. J. Anim. Sci. 46: 149-156.