Methane Production in Dairy Cows

P.W. MOE a n d H . F. T Y R R E L L

US Department of Agriculture Science and Education Administration Agricultural Research Animal Science Institute Ruminant Nutrition Laboratory Beltsville, MD 20705
ABSTRACT

The relationship among diet composition, intake, and methane production was investigated with data during 404 total energy balance trials with Holstein cows. Methane production in all trials was measured in open circuit respiration chambers. The most useful predictor of total methane production was amounts of soluble residue, hemicellulose, and cellulose that apparently were digested. The regression was methane (Meal) = .439 + .273 + .015 kg digestible soluble residue + .512 -+ .078 kg digestible hemicellulose + 1.393 + .097 kg digestible cellulose.
INTRODUCTION

diets. Our objective was to identify the relationship between dietary carbohydrate and methane production in cattle ingesting diets at a substantially greater range of intake than in the studies cited.

E X P E R I M E N T A L PROCEDURE

The systematic use of metabolizable energy (ME) to describe the energy of ruminant diets depends on either direct measurement of methane production or adequate means of estimating methane production. Because methane is a gaseous loss, its measurement requires specialized equipment. Therefore, the ME of many diets is estimated by calculated methane production. Kriss (7) found a linear relationship between methane production and dry matter intake of cattle. Bratzler and Forbes (2) with cattle and Swift et al. (14) with sheep identified a relationship between methane production and amount of carbohydrate apparently digested. Blaxter and Clapperton (1) concluded that methane production at maintenance by cattle and sheep could be described by the relationship: CHa (percentage of gross energy consumed) = 3.67 + .62 digestible energy (DE, %) and that the rate of change in percentage of gross energy lost as methane by an increase in intake equal to one times maintenance could be described by b = .054 DE(%) - 2.26 for mixed

Received March 28, 1979. 1979 J Dairy Sci 62:1583-1586

The data are from 404 total energy balance trials with Holstein cows in experiments designed to study the following effects on the energy of dairy cattle diets: percentage of protein (8); proportion of concentrate (15); relative energy value of corn grain and barley (17), oats (12), beet pulp (16), dried brewers grains, and dried distillers grains (18), or wheat bran (10); the physical form of corn grain in the total diet (9, 13); or the incremental energy value of corn grain (11). The experimental procedures and diets are described in the individual reports. Methane production was measured during three or four consecutive 24-h periods during each balance trial. An aliquot of air exhausting from the respiration chamber was collected continuously in a 10-liter spirometer (bell type) sealed with paraffin oil. Either two or three spirometers were used to collect replicate samples from each respiration chamber. The composition of chamber exhaust air collected in the spirometers was measured each day with an infrared analyzer previously calibrated with a reference gas mixture of known methane content. Methane energy was calculated from volume measurement by the factor 9.45 kcal/ liter (3). Chemical analyses of feeds and feces were based on the fiber methods o f Goering and Van Soest (5). These included acid-detergent fiber (ADF), cell walls (neutral-detergent fiber, NDF), cellulose (ADF-lignin-silica), and hemicellulose (NDF-ADF). An additional fraction, soluble residue, was calculated by subtracting crude protein and ether extract from the neutral-detergent solubles. The soluble residue

1583

1584

MOE AND TYRRELL the a m o u n t o f each c a r b o h y d r a t e c o m p o n e n t that apparently was digested as (R 2 = .73, Sy.x = .56): Ctt4 (Mcal/day) = .439 + .273 -+ .015 soluble residue (kg digested) + .512 -+ .078 hcmicellulose (kg digested) + 1.393 -+ .097 cellulose (kg digested) P r o d u c t i o n o f m e t h a n e per gram of cellulose digested is nearly three times that per gram of hemicellulose digested and five times that per gram of soluble residue digested. A part o f the latter difference may be due to a substantially lower p r o p o r t i o n o f soluble residue's being f e r m e n t e d in the rumen and a larger p r o p o r t i o n being digested and absorbed as glucose in the small intestine. A substantial a m o u n t of hemicellulose also m a y escape f e r m e n t a t i o n in the rumen. The p r o d u c t i o n of m e t h a n e f r o m 26 soluble carbohydrates was studied by Czerkawski and Breckenridge (4) in 53 experiments with an artificial rumen. T h e y c o n c l u d e d that, with the e x c e p t i o n o f r h a m n o s e , the a m o u n t of m e t h a n e p r o d u c e d was n o t depend e n t on the t y p e o f c a r b o h y d r a t e but rather the amount of c a r b o h y d r a t e f e r m e n t e d . The m e a n a m o u n t of m e t h a n e p r o d u c e d was 6 k c a l / 1 0 0 kcal of sugar f e r m e n t e d . If the gross energy value of c a r b o h y d r a t e a p p a r e n t l y digested in our e x p e r i m e n t is 4.2 kcal/g, the

fraction thus contains the m o s t soluble and readily f e r m e n t a b l e or digestible c a r b o h y d r a t e and for the m i x e d diets studied here likely consists primarily of starch. Statistical relationships were established by m u l t i p l e regression analysis with the least squares m e t h o d s of Harvey (6).
RESULTS A N D DISCUSSION

The mean, standard deviation, and range o f chemical c o m p o s i t i o n , intake of c a r b o h y d r a t e c o m p o n e n t s , and a m o u n t s apparently digested are in Table 1. The m a x i m u m a m o u n t o f total c a r b o h y d r a t e digested was 11.5 kg/day which is a p p r o x i m a t e l y twice that of the study of Bratzler and Forbes (2). A regression in which total m e t h a n e production was related to intake o f crude protein, ether extract, soluble residue, hemicellulose, cellulose, and lignin indicated nonsignificant terms for crude protein, ether extract, and lignin. A second regression related m e t h a n e p r o d u c t i o n to the remaining terms (R 2 = .67, S y.x = .62): CIt4 (Mcal/day) = .814 + .122 -+ .018 soluble residue (kg fed) + .415 + .074 hemicellulose (kg fed) + .633 + .076 cellulose (kg fed) Total m e t h a n e p r o d u c t i o n also was related to

TABLE 1. Mean and range of diet composition, intakes, and methane production. Variable Body weight Dry matter Crude protein Ether extract Soluble residue Hernicellulose Cellulose Lignin Protein Soluble residue Itemicellulose Cellulose Digestible soluble residue Digestible hemicellulose Digestible cellulose Methane production Methane production Units kg kg/day % % % % % % kg/day kg/day kg/day kg/day kg/day kg/day kg/day Mcal/day % of GE Mean 617 12.13 15.15 2.57 41.08 15.13 14.31 5.08 1.84 5.06 1.80 1.71 4.64 .86 .80 3.27 6.31 SD 101 4.28 2.53 .63 7.09 3.73 4.00 1.51 .73 2.10 .72 .68 1.91 .43 .36 1.07 1.46 Minimum 369 2.72 4.92 .90 18.08 7.07 8.53 2.16 .27 .95 .47 .41 .86 .01 .21 .91 1.60 Maximum 893 22.93 23.26 5.10 54.10 28.17 34.25 13.69 3.83 9.66 4.41 3.90 9.26 2.81 1.94 5.81 9.90

Journal of Dairy Science Vol. 62, No. 10, 1979

METHANE PRODUCTION TABLE 2. Effect of intake on methane production from digestible carbohydrate fractions. Mcal CH4 produced/kg carbohydrate digested Intake (X maintenance) N <1.5 1.5 - 2.5 2.5 - 3.5 >3.5 Intake >1.5 Alldata 103 119 152 30 301 404 SD .29 .48 .62 .81 .60 .56 R2 .66 .65 .61 .53 .65 .73 Intercept .462 .227 -.569 -.502 .322 .438 Soluble residue b .460 .285 .355 .360 .255 .273 SE b .040 .047 .047 .154 .023 .015 Hemicellulose b .538 .630 .515 .610 .451 .512 SE b .121 .153 .131 .421 .093 .078 b .634 1.521 1.957 1.372
1.699

1585

Cellulose SE b .107 .162 .170 .581 .119 .097

1.393

kcal m e t h a n e p r o d u c e d / 1 0 0 kcal o f carboh y d r a t e a p p a r e n t l y digested was 6.5 for soluble residue, 11.5 for h e m i c e l l u l o s e , a n d 33.6 for cellulose. O u r d a t a i n d i c a t e m a j o r d i f f e r e n c e s in the amount of methane produced during the digestion of various c a r b o h y d r a t e fractions. Experiments of the type described by Czerkawski a n d B r e c k e n r i d g e (4) are n e e d e d to establish r e l a t i o n s h i p s b e t w e e n f e r m e n t a t i o n of structural carbohydrates and methane production. The effect of intake on methane production was s t u d i e d b y deriving s e p a r a t e regressions w i t h i n four classes o f i n t a k e . T h e s e d a t a are in T a b l e 2. M e t h a n e p r o d u c t i o n was n o t i n f l u e n c e d as m u c h b y t y p e of c a r b o h y d r a t e digested a t low i n t a k e as at higher i n t a k e s . T h e differe n c e s a m o n g t h e partial regression c o e f f i c i e n t s for individual digestible c a r b o h y d r a t e f r a c t i o n s were less at i n t a k e s o f less t h a n 1.5 t i m e s m a i n t e n a n c e t h a n at h i g h e r i n t a k e s . M u c h o f this d i f f e r e n c e is associated w i t h t h e digestible

cellulose f r a c t i o n t h a t increased f r o m . 6 3 4 Mcal C H 4 / k g b e l o w 1.5 t i m e s m a i n t e n a n c e to 1 . 6 9 9 Mcal CI-14/kg a b o v e 1.5 t i m e s m a i n t e n a n c e . T h e l a t t e r rate of m e t h a n e p r o d u c t i o n is e q u i v a l e n t to 41% o f t h e cellulose e n e r g y a p p a r e n t l y digested. Likely, t h e high rate o f m e t h a n e p r o d u c t i o n a s s o c i a t e d w i t h cellulose d e g r a d a t i o n i n d i c a t e s a partial s h i f t f r o m a p r o p i o n a t e t y p e f c r m e n t a t i o n to m e t h a n o g e n i c f e r m e n t a t i o n in the rumen. The magnitude of methane product i o n associated w i t h cellulose digestion, h o w e v e r , suggests t h a t m e t h a n e w o u l d have to c o m e f r o m sources o t h e r t h a n t h e p r o d u c t s o f cellulose d e g r a d a t i o n a l o n e t o b a l a n c e t h e s t o i c h e o m e t r y o f t h e c o n v e r s i o n o f cellulose to e n d p r o d u c t s o f f a t t y acids a n d gases. P r o d u c t i o n o f m e t h a n e is i n f l u e n c e d b y t h e n a t u r e of t h e c a r b o h y d r a t e digested, b u t t h i s e f f e c t is relatively less i m p o r t a n t at low t h a n at high feed i n t a k e . In e f f o r t s to test t h i s o b s e r v a t i o n m o r e d i r e c t l y , regressions were derived for t h e d a t a w i t h i n several i n t a k e s ; t h e i n t a k e o f

TABLE 3. Effect of intake on error in prediction of methane production. Independent variable in prediction Intake (X maintenance) Dry matter intake (kg/day) Sy.x <1.5 1.5 - 2.5 2.5 - 3.5 >3.5 >1.5 103 119 152 30 301 .30 .62 .80 .87 .75 R2 .64 .40 .34 .40 .44 Digestible carbohydrate (kg/day) Sy,x .30 .64 .84 .89 .78 R2 .65 .36 .27 .37 .40 Digestible soluble residue, digestible hemicellulose, and digestible cellulose (kg/day) Sy. .29 .48 .62 .81 .60 R2 .66 .65 .61 .53 .65

Journal of Dairy Science Vol. 62, No. 10, 1979

1586

MOE AND TYRRELL Anim. Prod. Publ. No. 11:441. 4 Czerkawski, J. W., and G. Breckenridge. 1969. Fermentation of various soluble carbohydrates by rumen micro-organisms with particular reference to methane production. Brit. J. Nutr. 23:925. 5 Goering, H. K., and P. J. Van Soest. 1970. Forage fiber analysis: apparatus, reagents, procedures and some applications. 6 ttarvey, W. R. 1975. Least squares analysis of data with unequal subclass numbers. USDA ARS-H4. 7 Kriss, M. 1930. Quantitative relations of the dry matter of the food consumed, the heat production, the gaseous outgo and the insensible loss in body weight of cattle. J. Agr. Res. 40:283. 8 Moe, P. W., and H. F. TyrreIl. 1972. The net energy value for lactation of high- and low-protein diets containing corn silage. J. Dairy Sci. 55: 318. 9 Moe, P. W., and H. F. Tyrrell. 1977. Effects of feed intake and physical form on energy value of corn in timothy hay diets for lactating cows. J. Dairy Sci. 60: 752. 10 Moe, P. W., and H. F. Tyrrell. 1977. The incremental energy value of wheat bran for dairy cows. Northeastern Section ADSA-ASAS Mtgs. (Abstr.) 11 Moe, P. W., and |1. F. Tyrrell. 1979. Effect of endosperm type on incremental energy value of corn grain for dairy cattle. J. Dairy Sci. 62:447. 12 Moe, P. W., tl. F. Tyrrell, and N. W. Hooven, Jr. 1973. Energy balance measurements with corn meal and ground oats for lactating cows. J. Dairy Sci. 56:1149. 13 Moe, P. W., It. F. Tyrrell, and N. W. ltooven, Jr. 1973. Physical form and energy value of corn grain. J. Dairy Sci. 56:1298. 14 Swift, R. W., J. W. Bratzler, W. II. James, A. D. Tillman, and D. C. Meek. 1948. The effect of dietary fat on utilization of the energy and protein of rations by sheep. J. Anita. Sci. 7:475. 15 Tyrrell, H. F., and P. W. Moe. 1972. Net energy value for lactation of a high and low concentrate ration containing corn silage. J. Dairy Sci. 55:1106. 16 Tyrrell, tl. F., P. W. Moe, and L. S. Bull. 1973. Energy value of cracked corn and dried beet pulp fed to Flolstein cows. J. Dairy Sci. 56:1384 (Abstr.) 17 Tyrrell, tt. F., and P. W. Moe. 1974. Net energy value for lactation of a corn and a barley ration. J. Dairy Sci. 57:451. 18 Tyrrell, H. F., and P. W. Moe. 1979. Unpublished data.

d r y m a t t e r , t o t a l digestible c a r b o h y d r a t e , and individual c a r b o h y d r a t e c o m p o n e n t s were indep e n d e n t variables. The s t a n d a r d errors and R 2,s are in Table 3. A t intakes b e l o w 1.5 t i m e s m a i n t e n a n c e , m e t h a n e p r o d u c t i o n is nearly as related to e i t h e r d r y m a t t e r intake or total digestible c a r b o h y d r a t e alone as t o individual digestible c a r b o h y d r a t e c o m p o n e n t s (R 2 = .64, .65, and .66). A t intakes a b o v e 1.5 times maintenance, however, consideration of the individual a m o u n t s o f soluble residue, hemicellulose, and cellulose a p p a r e n t l y digested p r o v i d e d a s u b s t a n t i a l i m p r o v e m e n t in the p r e c i s i o n with w h i c h m e t h a n e p r o d u c t i o n was p r e d i c t e d (R z = .65 v s . . 4 4 and .40) as c o m p a r e d to e i t h e r dry m a t t e r or total c a r b o h y d r a t e digested. S e p a r a t e regressions were o b t a i n e d for diets b a s e d o n corn silage and t h o s e c o n t a i n i n g hay. No d i f f e r e n c e s were a p p a r e n t , and we c o n c l u d e that the carbohydrate c o m p o n e n t s of these forage classes i n f l u e n c e m e t h a n e p r o d u c t i o n similarly. We c o n c l u d e t h a t w h e r e a s the m e t h a n e p r o d u c t i o n b y a d u l t cattle at m a i n t e n a n c e can be p r e d i c t e d a d e q u a t e l y f r o m dry m a t t e r or total digestible c a r b o h y d r a t e , a c c u r a t e prediction o f m e t h a n e p r o d u c t i o n by lactating dairy cattle requires d e t e r m i n a t i o n o f the n a t u r e o f the c a r b o h y d r a t e as well.

REFERENCES

1 Blaxter, K. L., anti J. L. Clapperton. 1965. Prediction of the amount of methane produced by ruminants. Brit. J. Nutr. 19:511. 2 Bratzler, J. W., and E. B. Forbes. 1940. The estimation of methane production by cattle. J. Nutr. 19:611. 3 Brouwer, E. 1965. Report of sub-committee on constants and factors. Proc. 3rd Sympos. on Energy Metabolism, Troon, Scotland, Europ. Ass.

Journal of Dairy Science Vol. 62, No. 10, 1979