Material and methods Directory UMM :Data Elmu:jurnal:L:Livestock Production Science:Vol64.Issue2-3.Jun2000:

254 T . Yan et al. Livestock Production Science 64 2000 253 –263

1. Introduction major differences due to level of intake. Over the last

2 decades cow genetic merit has substantially in- creased Coffey, 1992 and these animals can The metabolisable energy ME concentration in produce milk yields of over 50 kg day and consume the diet of ruminant animals is determined as the more than 300 MJ day of ME Yan et al. 1997a, a difference between gross energy GE intake and level which can be over five times their maintenance energy outputs in faeces, urine and methane. The requirement as estimated from the Agricultural and measurement of methane energy output CH -E 4 Food Research Council 1990. Beever et al. 1998 requires complex and expensive equipment and 0.75 hence prediction equations are widely used to calcu- also reported a mean ME intake of 1.89 MJ kg late CH -E. A number of these equations have been with lactating cows, which is approximately four 4 published since 1930, using total dry matter DM levels of feeding calculated from the Agricultural intake Kriss, 1930; Axelsson, 1949, digestible and Food Research Council 1990. The effect of carbohydrates Bratzler and Forbes, 1940; Moe and feeding level on methane production is therefore Tyrrell, 1979, energy digestibility and feeding level becoming of increasing importance and there is Blaxter and Clapperton, 1965, and a range of therefore a need to re-examine this effect. animal and dietary factors Holter and Young, 1992. At this Institute, between 1992 and 1997 a number However, the data used to develop these equations of experiments have been completed with lactating were collected from ruminant animals offered diets dairy cows n 5247 and beef steers n575 offered containing mainly dry or high DM forages, rather grass silage-based diets and subjected to gaseous than low DM grass silages, typical of those used in exchange measurements in calorimetric chambers. many areas of Western Europe. The fermentation The objective of the present study was therefore to process in low DM grass silage results in a low use the energy metabolism data from these studies to concentration of water soluble carbohydrates and evaluate the relationship between methane product- high levels of fermentation products, such as volatile ion and a number of animal and dietary factors. fatty acids VFAs, lactate, alcohol and ammonia. Furthermore, the concentration of fibre in the DM of silage can also differ from that in dried grass due to

2. Material and methods

the ensiling process and differences in harvesting date. The feeding of grass silage can therefore result 2.1. Animals and calorimeters in different fermentation patterns in the rumen when compared to that of dried forages. This effect can A total of 322 cattle including 247 lactating dairy shift the proportion of acetic acid in total VFAs cows and 75 beef steers were subjected to measure- produced in the rumen, resulting in change of ments of energy metabolism in calorimetric cham- methane production Ørskov and Ryle, 1990. An- bers at the Agricultural Research Institute of North- derson and Jackson 1971 reported a consistently ern Ireland between 1992 and 1997. The dairy cattle higher proportion of acetic acid in total VFAs in the were Holstein Friesian cows and the beef cattle were rumen of sheep offered grass silage rather than grass Charolais cross, Simmental cross and Aberdeen hay. The CH -E was found to account for a higher Angus cross steers. The dairy cows were drawn from 4 proportion over GE intake or digestible energy DE ten feeding experiments Cushnahan et al., 1995; intake in dairy cattle offered diets based on grass Gordon et al., 1995a,b, 2000; Carrick et al., 1996; silages, rather than dried grass Yan et al., 1997b. Yan et al., 1996, 1997b; Keady and Mayne, 1998; As a consequence, the prediction of CH -E output Ferris et al., 1999; C.S. Mayne, personal communi- 4 with grass silage-based diets using the equations cation, and the beef cattle were obtained from 3 developed from the non-grass silage-based diets, feeding experiments Kirkpatrick, 1995; Kirkpatrick may result in considerable error in the calculation of et al., 1997; Lavery, 1998. In each experiment the ME intake for animals. Therefore, there is a need to animals were offered the experimental diets for at examine these effects with grass silage-based diets. least 3 weeks in individual feeding accommodation In addition to differences in diet, there are also before measurement of energy metabolism. In the T . Yan et al. Livestock Production Science 64 2000 253 –263 255 metabolism unit, each animal was subjected to a Thirty-five dairy cows and twelve beef steers each 6-day diet balance measurement with total faeces and from a single experiment were offered silage as the urine outputs being collected. Immediately after sole diet, but otherwise in all other experiments the completion of the balance measurement, each animal dairy and beef cattle n 5275 were offered the was transferred to respiration calorimeters. The silages with a range of proportions of concentrates animals remained in the calorimeters for 3 days with from 0.146 to 0.815 DM basis with a mean of measurement of gaseous exchange over the final 0.467 S.D. 0.1484. The concentrates used in each 48-h period. The dairy cows were of various genetic of the studies included a mineral vitamin supplement merits and at a range of lactation stages milk yield and some of the following ingredients, from 3.2 to 49.1 kg day with mean of 23.2 S.D. 8.07 kg day. The lactation number for the dairy Cereal grains: Barley, wheat or maize cows ranged from 1 to 9 and liveweight from 416 to By-products: Maize gluten meal, molassed sugar-beet 733 mean 565, S.D. 23.6 kg. The age and pulp, citrus pulp or molasses liveweight of the beef cattle ranged, respectively Protein supplements: Fish meal or soya-bean meal from 18 to 21 months and from 450 to 644 mean 531, S.D. 31.1 kg. The concentrate portion of the diet was offered The calorimeters used in the present study were either in a complete diet mixed with the grass silage, indirect open-circuit respiration calorimeters. All or as a separate feed from the silage. All animals equipment, procedures, analytical methods and were offered either silage or the complete diet ad calculations used in the calorimetric experiments libitum. The data on mean, S.D. and range for were as reported by Gordon et al. 1995b. Cali- animal, silage composition, total diet and energy bration of the chambers is carried out in two stages, utilisation variables are presented in Table 1. As the i.e. analyser calibration and flow calibration. Firstly, data set is combined from both lactating dairy cows the analysers are calibrated with gases produced and beef steers, there was a wide range in live from individual pure analytical standard gases weight, DM intake, feeding level, methane energy methane, carbon dioxide and oxygen using Wostoff output and energy intake. Mixing pumps. This determines the absolute range Gross energy GE concentration in silages was 0–500 ppm for methane and the linearity within determined using undried silages in an adiabatic this range. Before each run the analysers are cali- bomb calorimeter Gallenkamp, Loughborough, UK brated using oxygen free nitrogen and a gas of Porter, 1992. Silage DM concentration was de- known concentration span gas. This calibration is termined on an alcohol–toluene basis, which was checked automatically every 6 h. Secondly, the flow subsequently used as a basis of expressing all measurement systems are checked with analytical nutrient concentrations in silages. Analysis for other grade carbon dioxide and nitrogen using, respective- nutrients in feeds, faeces and urine were as described ly the carbon dioxide and oxygen analysers, by by Gordon et al. 1995b and Mayne and Gordon determining the recovery of carbon dioxide and 1984. depletion of oxygen. 2.3. Data analysis 2.2. Diets The relationship between CH -E and energy in- 4 A total of 30 perennial ryegrass silages were take and or other variables was examined on the examined over the 13 experiments. The silages combined data of dairy and beef cattle in four steps. encompassed primary growth and first and second Firstly, CH -E was related to total GE intake GEI 4 regrowth material. The grass was either unwilted or and digestible energy DE intake DEI using the wilted prior to ensiling and ensiled with or without linear regression technique Eq. I. Secondly, CH - 4 application of silage additives. All silages were well E was calculated as CH -E GEI and CH -E DEI. 4 4 preserved with silage dry matter DM concentration These two variables were then each related to FL, at feeding ranging from 168 to 398 g kg Table 1. apparent energy digestibility, silage DM intake 256 T . Yan et al. Livestock Production Science 64 2000 253 –263 Table 1 Data on animal, diet and energy metabolism for both dairy and beef cattle Mean S.D. Minimum Maximum Cattle n 5322 Live weight kg 563 58.9 416 733 a Milk yield kg day 23.2 8.07 3.2 49.1 Silage composition n 530 Dry matter g kg 223 55.1 168 398 pH 3.95 0.336 3.60 4.75 NH -N total-N g kg 91 60.3 22 240 3 Crude protein g kg DM 145 26.8 97 193 Gross energy MJ kg DM 18.7 0.53 17.8 20.0 Ash g kg DM 87 13.6 66 117 Acid detergent fibre g kg DM 375 53.9 301 486 Total diet n 5322 DM intake kg day 14.61 4.939 5.55 24.26 Silage DM total DM intake 0.544 0.2334 0.169 1.000 Total ADF total DM intake 0.248 0.0615 0.137 0.372 Silage ADF total ADF 0.782 0.1538 0.409 1.000 Feeding level 3.23 1.061 1.28 5.71 Agricultural and Food Research Council, 1990 Energy utilisation MJ day n 5322 Gross energy intake 271.9 92.34 105.5 454.6 Digestible energy intake 207.0 70.58 79.5 351.6 Metabolisable energy intake 177.4 61.81 66.1 311.2 Methane energy output 18.1 5.88 4.1 29.6 a For dairy cows only n 5247. S as a proportion of total DM intake T CH -E 5 a 1 b ? intake 1 c ? [FL-1] IIIa DMI DMI 4 S T , total ADF intake T as a propor- DMI DMI ADFI tion of T T T , and silage ADF intake CH -E 5 a 1 intake ? b 1 c ? [dietary factor] IIIb DMI ADFI DMI 4 S as a proportion of T S T Eqs. ADFI ADFI ADFI ADFI IIa–c. Thirdly, CH -E was related to energy intake CH -E 5 a 1 intake ? b 1 c ? [dietary factor] 4 4 GE or DE and FL above maintenance FL-1 or 1 d FL-1 IV dietary factor T T , S T or S ADFI DMI DMI DMI ADFI T Eqs. IIIa–b. Finally, CH -E was predicted ADFI 4 The above equations were fitted, respectively to using the above three groups of variables, i.e. energy the following three equations intake GE or DE, feeding level FL-1 and dietary factor T T , S T or S T y 5 a 1 b1 ? x1 ADFI DMI DMI DMI ADFI ADFI i Eq. IV y 5 a 1 b1 ? x1 1 b2 ? x2 i y 5 a 1 b1 ? x1 1 b2 ? x2 1 b3 ? x3 i CH -E 5 a 1 b ? intake I 4 where a represents the effect of experiment i for i i51 to 13, x1, x2, and x3 are the x-variables and b1, CH -E intake 5 a 1 b ? digestibility IIa 4 b2 and b3 are their regression coefficients. The feeding level FL is estimated as multiples of ME CH -E intake 5 a 1 b ? [FL-1] IIb 4 intake MEI over ME requirement for maintenance ME MEI ME , where ME was calculated m m m CH -E intake 5 a 1 b ? [dietary factor] IIc from the equations of the Agricultural and Food 4 T . Yan et al. Livestock Production Science 64 2000 253 –263 257 Research Council 1990. The statistical programme ing from 0.639 to 0.847 had no significant relation- used was GENSTAT 5 Genstat 5 Committee, 1993. ship with either CH -E GEI or CH -E DEI when 4 4 using Eq. IIa. However, when using Eqs. IIb–c feeding level and dietary factors were each sig-

3. Results nificantly P ,0.001 related to CH -E GEI or CH -