effect of sodium bicarbonate on nitrogen utilization and

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EFFECT OF SODIUM BICARBONATE ON NITROGEN UTILIZATION ANDFEED INTAKE BY LAMBSL. E. PHILLIP

Department of Animal Science, Macdonald Carnpus of McGill University, Ste.Anne de Bellevue, Quebec HgX IC). Received l3 Jan. 1983, accepted 2 May1983.Psrr-lrp, L. E. 1983. Effect of sodium bicarbonate on nitrogen utilization andfeed intake by lambs. Can. J. Anim. Sci. 63: 613-62 l.Eight wether lambs, four equipped with rumen cannulae and four intact, wereassigned to four high-concentrate diets according to a 4x4 double latin squarewith 27-day periods. The basal diet consisted of barley and corn silage, the latterrepresenting 3OVo of the dry matter (DM), and was either unsupplemented (97cdietary protein) or supplemented (157c protein) with brewers'dried grains. Toeach of the low protein (LP) and high protein (HP) diets, sodium bicarbonate(NaHCO.) was either not added or added at the level of 4Vo of the diet DM. Therewas a marked, though not statistically significant, improvement in nitrogen reten-tion due to NaHCO, inclusion, particularly when protein intake was low. Voluntaryfeed intake increased by ll%o as a result of NaHCO. addition but the effect wasnonsignificant. Bicarbonate-induced changes in feed intake were unrelated tochanges in rumen pH. A maximum ruminal osmolality of 303 mOsm/kg elicitedby NaHCO, addition did not appear to restrict feed intake. Rumen ammonla con-centration tended to decrease as a result of NaHCO. addition. Digestibility oforganic matter and of DM was unaffected by the inclusion of NaHCO, but diges-tibility of nitrogen tended to increase. It is suggested that NaHCO, could have aprotein-sparing effect when added to protein-deficient concentrate diets, and thatbicarbonate-induced changes in feed intake may be related to its influence onprotein status of ruminants.Key words: Bicarbonate, buffers, lambs, nitrogen utilization, feed intakelEffet du bicarbonate de sodium sur I'assimilation de I'azote et l'indice de con-sommation des aliments chez l'agneau.]Titre abr6g6: Bicarbonate de sodium pour agneaux.Quatre rations trds concentr6es ont 6t6 servies pendant 27 jours ir huit agneauxchAtrds, dont la moiti6 avait une fistule dans le rumen, selon un carr6 latin doublede 4 x 4. La ration 6l6mentaire compos6e d'orge et d'ensilage de mais, ce dernierrepr6sentant 30Vo


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614 cANADTAN JouRNAL oF ANTMAL scrENcEl'azote a sembl6 s'am6liorer. On pense que l'addition de NaHCO, aux concentrdspauvres en prot6ines pourrait produire un effet d'6pargne sur 1'azote et que lesmodifications dans la prise alimentaire pourraient 6tre li6es ir la r6tention de I'azotechez les ruminants.Mots cl6s: Bicarbonate. tampon, agneau, utilisation de I'azote, consommationd'aliments

Sodium bicarbonate (NaHCO.) is used asan additive to grain- or silage-based dietsto offset the deleterious nutritional effectsof low ruminal pH and to aid ruminants intheir homeostatic adjustments to an acidIoad. Trenkle (1979) has discussed the pos-sible growth-limiting effect of metabolicacidosis in ruminants and implied that so-dium bicarbonate could have a sparing ef-fect on muscle protein catabolism. How-ever. the interaction between bicarbonateand protein metabolism in ruminants hasnot been fully explored.Asplund et al . (1980) reported an im-provement in nitrogen baiance with l7o so-dium bicarbonate added to a concentratediet but observed no significant interactionwith protein source (soybean meal or corngluten meal). The results of their growthtrial, however, revealed that sodium bicar-bonate caused a greater increase in weightgain at 9Vo dietary protein than at 127o pro-tein, indicating that bicarbonate could, infact, have a protein-sparing effect in thenutrition of ruminants. Huntington et al.(1911) reported, on the other hand, that theaddition of 27o sodium bicarbonate to aprotein deficient diet did not affect growthrate of lambs. It appears, therefore, thatthe effect of sodium bicarbonate on nitro-gen utilization by lambs is unclear.The addition of bicarbonate to concen-trate diets also enhances feed intake but themechanism by which this response occurshas not been clearly defined. Trenkle(1979) has related bicarbonate-inducedchanges in feed intake to an increase inrumen pH. However, not only is the effectof bicarbonate on rumen pH equivocal(Bush et al. 1981; Nicholson et al. 1963a;Farhan and Thomas 1978), but Kellawayet al. (1978) observed no significant effectof bicarbonate on rumen pH despite marked

increases in voluntary feed intake by calvesfed a concentrate diet (Kellaway et al.1977). Egan and Moir (1965) have sug-gested that changes in feed intake by rum-inants could be mediated through changesin protein status of the animal. It is, there-fore, possible that the effect of sodium bi-carbonate on feed intake is related to itsinfluence on nitrogen metabolism of rum-inants.The objective of this study was to testthe hypothesis that NaHCO. enhances ni-trogen utilization by lambs consuming alimited amount of protein (relative to re-quirement), and that bicarbonate-inducedchanges in feed intake are associated withcorresponding changes in nitrogen reten-tlon.MATERIALS AND METHODSAnimal Management and DietsThe experiment was conducted with eightwether lambs (Suffolk crossbred) weighing20+ 1.1 kg. Four of the iambs were equippedwith rumen cannulae. All eight lambs wereplaced in individual pens in an air-conditionedroom maintained at a temperature of 20'C anda photoperiod regime of 10 h light:l4 h dark.The pens were bedded with wood shavings.Prior to the start of the trial. the lambs weretreated for internal parasites with RipercoiR so-lution (Cyanamid Canada Inc., Montreal, Que.)and injected subcutaneously with Covexin-8R(Bunoughs-Wellome, Montreal). Each animalreceived an intramuscular injection of 1 mL vi-tamin mixture to provide 500 000 IU. vitaminAr, 75 000 IU, vitamin D,, and 50 IU vitaminE. Trace-mineralized salt blocks were providedto all animals. The lambs had ad libitum accessto fresh water and were offered the experimen-tal diets in two equal portions at 0900 h and at1700 h.The basal diet consisted of barley and cornsilage, the Iatter representing 3OVo of the dietdry matter (DM). Brewer's dried grains (BDG)were used as a source of supplemental protein

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PHILLIP SODIUM BICARBONATE FOR LAMBS 615because of their high potential for promotingnitrogen deposition in ruminants (Merchen etal. 1979). The calcium:phosphorus ratio of alldiets was maintained aI 2:l by the addition oflimestone. The experimental diets were for-mulated to provide either 157o or l\Vo dietaryprotein (no protein supplement) with or without47c NaHCO,. This level of bicarbonate waschosen because several studies have revealedthat when added at levels between 2Vo and 6Vc.NaHCO. generally improves voluntary intake ofconcentrate diets (Trenkle 1979). The sodiumbicarbonate was mechanically mixed with thesupplemental ingredients and at feeding time,the supplement mixture was combined withcorn silage, and hand-mixed.Experimental Design and MeasurementsEach group of lambs (four intact and four ru-men-fistulated) was assigned randomly to thefour experimental diets according to a 4 x 4double latin square with 27-day experimentalperiods. Voluntary intake was determined dur-ing the first 17 days, the last 7 days of whichrepresented the measurement phase. The lambswere then transferred to metabolism cages andduring the remaining 10 days, nitrogen balancewas measured. The lambs were allowed a 5-dayperiod of adaptation prior to the 5-day collec-tion of urine and feces. To estimate voluntaryintake, the lambs were offered enough feed toensure 15% feed refusal (orts). During the 7-day measurement phase, the orts were collecteddaily and stored at 5"C. At the end of this periodthe orts were composited and stored at - lO"Cfor subsequent analysis. Samples of the supple-ment mixture and of the corn silage were alsotaken daily. The corn silage was stored at- 10"C for subsequent analysis.For the determination of nitrogen balance thelambs were fed 75 g DM/Wk,O i5 and fitted withplastic bags for feces collection. The bags wereheld in place with rubber cement. Urine wascollected in polyethylene bottles containing 50mL 10 N HrSOo as previously described (Phillipand Buchanan-Smith 1982). Daily samples ofdiluted urine were stored at 5'C. At the end ofthe 5-day period, the individual urine samplesfrom each lamb were pooled and a subsamplewas stored at - l0"C for subsequent nitrogenanalysis. Samples of feed and orts (if any) weretreated in the same manner as was described forsamples obtained during the voluntary intakemeasurements.

On the last day of the collection period rumenfluid was withdrawn from the four fistulatedlambs with the use of a vacuum pump. The sam-ples were withdrawn 10 min before feeding andagain at 0.5, 1.0, 2.0, 4.0 and 6.0 h after feed-ing. The pH of the rumen fluid samples wasdetermined immediately. The rumen fluid wasthen strained through four layers of cheeseclothand centrifuged at 10 000 g for 15 min. A por-tion of the supernatant was immediately storedat 10'C for subsequent determination of am-monia. No preservative was added. Osmolalitywas determined on an aliouot of the fresh su-pernatant.Analytical MethodsDry mattel' content of corn silage and of feceswas determined after the samples had beendried in a forced-draught oven at 50'C for 48h. Ash and nitrogen content of the dried sam-ples and the nitrogen content of urine were ana-lyzed according to the methods of the Associ-ation of Official Agricultural Chemists (19'75).Acid detergent fiber (ADF) was determined ac-cording to the procedure of Van Soest (1963).Ammonia-nitrogen was determined on a diluted(10-fold) sample of rumen fluid with the use ofthe ammonia electrode (model 95-10, OrionResearch Inc., Cambridge, Mass.) connected toa pH-ion meter (Acumet 750, Fisher Scientific).Osmolality was measured with an automaticosmometer (Osmette-Model S, Precision Sys-tems Inc., Sudbury. Mass.).Analysis of variance of the data was per-formed by means of the general linear modelprocedure of the Statistical Analysis System(SAS Institute Inc., Box 8000, Cary, NorthCarolina 27511 .). One fistulated lamb sufferedillness after I mo on trial and was removed fromthe experiment.

RESULTS AND DISCUSSIONThe basal diet of corn silage and barleygrain contained l0.7Vo protein (Table l).Supplementation with BDG increased theprotein content slightly above the calcu-lated value of 15Vo. The inclusion of BDGalso resulted in a slight increase in ADFcontent. Addition of sodium bicarbonatecaused an expected increase in the ash con-tent of both the high protein (HP) and thelow protein tLPl diets.

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PHTLLIP SODIUM BICARtsONATE FOR LAMBS 617hand. showed that when weaned calveswere fed a concentrate diet contaimng 47oor 7olo NaHCOr, nitrogen balance was im-proved. As indicated earlier, it is likelythat sodium bicarbonate influences nitro-gen balance only when dietary protein islimiting. This condition probably occursmore often in young animals than in semi-mature or mature animals.Although nitrogen digestibility was im-proved with the addition of NaHCO. (Table3), the improvement in nitrogen balance,particularly with the LP diet, was duemainly to a reduction in urinary nitrogenexcretion (Table 2). The 207o increase innitrogen digestibility as protein level in-creased from 10 to l5Vo is consistent withthe general increase in nitrogen digestibil-ity with increasing dietary protein, an ob-servation which has also been made withforages (Egan and Kellaway 1971). Diges-tibility of organic matter and of dry matter(Table 3) was unaffected (P>0.10) eitherby protein level or by the addition ofNaHCO.. The interaction was also nonsig-nificant. It appears that there is no im-provement in energy digestibility by 20-kglambs when dietary protein levels are 107oor more. In their studies with steers Ni-cholson et al. (1963a,b) found no effecfofsodium bicarbonate on organic matter di-gestibility. The addition of up Io JVcNaHCO. to a barley-based diet also failedto alter dry matter digestibility by calves(Leibholz et al. 1980). In studies withsheep fed grass silages, the inclusion ofhish levels of sodium bicarbonate also had

no significant effect on either dry matteror organic matter digestibility (Farhan andThomas 1978).Feed Intake and Rumen ParametersVoluntary intake of dry matter and of or-ganic matter (OM) was enhanced as a re-sult of sodium bicarbonate addition (Table3) but the effect was not statistically sig-nificant (P>0.10). This lack of signifi-cance could be explained by the high de-gree of variability in feed consumption, asindicated by the magnitude of the standarderrors. The interaction between sodium bi-carbonate and protein level on feed intakewas also nonsignificant. In their earlierwork with steers fed an all-concentrate(barley-based) diet, Nicholson et al.(1963a) found that when allowances weremade for the presence of 37o NaHCO. inthe diet, the addition of the buffer signifi-cantly improved feed consumption. Hun-tington et al. (1977) also observed in-creases in food intake by lambs fed a highconcentrate diet (907c ground corn) con-taining etther 2c/o or 4o/o NaHCO.. How-ever, in a recent study with lambs fed bar-ley-based diets, Bush et al. (1981) reporteda 10Vc reduction in feed intake when 2%NaHCO. was added to the diet. Althoughbarley constituted the major part of the ra-tion in this study, corn silage formed 307oof the dry matter. It is not known whetherthis dietary modification is related to theobserved effects of bicarbonate on feed in-take by these lambs.Although NaHCO. addition appeared to

Table 3, Effects of sodium bicarbonate and protein level on daily voluntary intake and digestibility by lambsLow protein High protein

Dry matter (DM) intake(g/w*.ntt)Organic rnatter (OM), intake(g/w*rnt')DM digestibility (7c)OM digestibility (6/c)Nitrogen digestibility (%)

72.8 + 5.83770.6 + 5.64

81 .5 + 6.0078.1 +5.8270.4 + 1 .4171.7+1.236r.4 !2.40

70. I + 1.37l2.l + |.1958.0 + 2.33

80.615.u3 90.6+5.8377 .1 + 5.64 86.6 i 5.6468.8 :! 1 .37 69 .0 + 1 .3170.6 + 1.19 70.3 1 1.1912.6+2.33 74.2+2.33

iValue: are merns: standard errors of means

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618 CANADIAN JOURNAL OF ANIMAL SCIENCEincrease rumen pH (Fig. 1), the bicarbon-ate-induced changes in voluntary feed in-fake were unrelated to changes in rumenpH. At 2 h after feeding, for instance, bi-carbonate addition increased rumen pH by0.23 units (6.20-6.43) and increased OMintake by llTo (Table 3). There was alarger change in rumen pH for the HP dietbut the extent of improvement in OM in-take was smaller (9Vo). The lack of anyconsistency between voluntary feed intakeand changes in rumen pH agrees with theresults of Kellaway et al. (1978). RumenpH showed the usual decline as timeelapsed after feeding. The changes in ru-men pH and other rumen parameters (am-monia, Fig. 2 and osmolality, Fig. 3) werenot statistically analyzed because, at pres-ent, there seems to be no appropriate modelto allow meaningful interpretation of thefrequent interactions on rumen measure-ments between treatment and rumen sam-pling time. Average values were also notcomputed for reasons discussed by Mahad-evan et al. (1982).Despite the slight elevation in rumen os-

molality with the addition of NaHCO.(Fig. 3) to the HP diet, feed intake was notdepressed. It should be noted though, thatthe changes in ruminal osmolality as timeafter feeding elapsed were relatively smalland inconsistent. After infusing osmoti-cally active compounds into the rumen ofsheep, Phillip et al. (1981) reported a lin-ear reduction in short-term intake of feedas rumen osmolality increased from 200 to550 mOsm/kg, a range that encompassesthe maximum value of 303 mOsm/kg(HP + NaHCO. diet) recorded in this study.The present finding that rumen osmolalitydid not limit the intake response to bicar-bonate is consistent, nevertheless, with thesuggestion of Bergen (1912) that below400 mOsm/kg rumen osmolality does notlimit feed intake by sheep.When NaHCO. was added to the HP dietthere was a marked decrease in rumen am-monia (Fig. 2). However, there was essen-tially no difference in rumen ammonia withand without bicarbonate addition to the LPdiet. In the absence of NaHCO., the dif-ference in rumen ammonia between the LP

d

0

Ti meFig. 1. Changes in rumen pH as aft'ected by

L24af ter feed'tng (h )protein level and sodium bicarbonate.

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PHILLIP - SODIUM BICARBONATE FOR LAMBS

Low protein (LHigh protein (HP)LP + llaHC03HP + NaHC0,

6t9

_ 16O

12

=9R O

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CANADIAN JOURNAL OF ANIMAL SCIENCEand the HP diets was 8 . 6 mg/ 1 00 mL ( I 0. 2vs. 18.8) at 0.5 h after feeding. WhenNaHCO: was included, the difference nar-rowed during the same interval to 3.4 mgNH3-N/100 mL (1 1.9 vs. 15.3) indicating,perhaps, that in the presence of NaHCO.,microbial utilization of rumen ammoniamay have been improved. If the reductionin rumen ammonia as a result of NaHCO.addition were truly a reflection of in-creased microbial growth resulting in anincreased supply of amino acids to the duo-denum (Harrison and McAllan 1980), thenthe effect of NaHCO, in improving feedintake could also have been mediatedthrough its influence on protein status ofthe lambs (Egan and Moir 1965).This study does not establish whether theimprovement in protein status (nltrogen re-tention) elicited with NaHCO. was the re-sult of increased duodenal flow of rumenmicrobial protein or the result of a reduc-tion in urinary excretion of ammonia(Trenkle 1979). No measurements weremade of the acid-base status of the lambsand thus it is difficult to invoke the conceptthat bicarbonate was producing the effectby minimizing urinary excretion of nitro-gen in animals that were "acid stressed"(Austic and Calvert l98l). On the otherhand, although sodium bicarbonate im-proves the efficiency of synthesis of mi-crobial protein from ammonia (Harrisonand McAllan 1980), it is still possible thatthe reduced levels of rumen ammonia werea reflection of an increased rate ammoniaabsorption at elevated rumen pH. Further-more. the effect of bicarbonate on rumenammonia levels is still a controversial issue(Okeke et al. 1983; Haaland and Tyrrell1982). The absence of the usual increasein rumen ammonia shortly after I'eeding theHP diet (Fig. 2) emphasizes the compli-cations in interprcting changes in rumenammonia in relation to nitrogen utilizationin ruminants.More studies may be needed to clarifythe precise mode of action of bicarbonateand its possible nitrogen-sparing effect at

low intakes of protein. It appears, never-theless. that if sodium bicarbonate is addedto concentrate diets for growing ruminants.it may be possible to reduce the dietaryprotein allowance without sacrificing ani-mal performance. This idea is consistentwith the report of Asplund et al. (1980)who found that sodium bicarbonate im-proved growth rate of lambs fed a concen-trate diet containing only 9Vo dietary pro-te1n.

ACKNOWLEDGMENTSThe author acknowledges the assistance of Mrs.Theresa Clark in managing the lambs and theadvice of Dr. J. F. Hayes in statistical analysisof the data. Technical assistance provided byMr. Jos6 Carreiio is gratefully appreciated. Thiswork was supported by a research operatinggrant from Agriculture Canada.ASSOCIATION OF OFFICIAL AGRICUL-TURAL CHEMISTS. 1975. Official methodsof analysis, l2th ed. AOAC, Washington, D.C.ASPLUND. J. M., GREBING, S. E., LEWIS,O . BOWMAN. J. and JENSEN, D. O. 1980.Interaction of buffer and protein solubility insheep diets. 72nd Ann. Meeting of the Ameri-can Society of Animal Science, Cornell Uni-versiry. J. Anim. Sci. 5l: (Suppl. l): 342.AUSTIC, R. E. and CALVERT, C. C. i981.Nutritional interrelationships of electrolytes andamino acids. Fed. Proc. 40: 63-6'7.BUSH, R. S.. NICHOLSON, J. W. G., Mc-INTYRE, T. M. and CALDER, F. W. 1981.The effect of sodium bicarbonate, cement kilndust and ammonium perchlorate on growth oflambs. Can. J. Anim. Sci. 61: 423-428.BERGEN, W. G. 19'72. Rumen osmolality asa factor in feed intake control of sheep. J.Anim. Sci. 34: 1054-1060.EGAN. A. R. and KELLAWAY, R. C. 1971.Evaluation of nitrogen metabolites as indices ofnitrogen utilization in sheep given frozen anddry mature herbages. Br. J. Nutr. 26:335-351.EGAN, A. R. and MOIR, R. J. 1965. Nutri-tional status and intake regulation in sheep. lEffects of duodenally infused single doses ofcasein, urea and propionate upon voluntary in-take of low protein roughage by sheep. Aust.J. Agric. Res. 16: 431 449FARHAN. S. M. and THOMAS, P. C. 1978.The effect of oartial neutralization of formic

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PHILLIP SODIUM BICARBONA'TE FOR LAMBS 621acid silages with sodium bicarbonate on theirvoluntary intake by cattle and sheep. J. Br.Grassland Soc. 33: l5l-158.HAALAND, G. L. andTYRRELL. H. F. 1982.EfTects of limestone and sodium bicarbonatebuffers on rumen measurements and rate of pas-sage in cattle. J. Anim. Sci. 55: 935-942.HARRISON, D. G. and McALLAN. A. B.1980. Factors affecting microbial growth yieldsin the reticulo-rumen. Pages 205-226 In Y.Ruckebusch and P. Thivend, eds. Digestivephysiology and metabolism in ruminants. AVIPublishing Co. Inc., Westport, Conn.HUNTINGTON, c. B., EMERICK, R. J. andEMBRY, L. B. 1977. Sodium bentonire or so-dium bicarbonate as aids in feeding high-con-centrate diets to lambs. J. Anim. Sci.45:804-8 1 1 .KELLAWAY, R. C., THOMSON, D, J.,BEEVER, D. E. and OSBOURN, D. F. 1977.Effects of NaCl and NaHCO, on food intake,growth rate and acid base balance in calves. J.Agric. Sci. (Camb.) 88: l-9.KELLAWAY, R. C., BEEVER. D. E,.THOMPSON, D, J., AUSTIN, A. R., CAM-MELL, S. B. and ELDERFIELD. M. L. 1978.The effect of NaCl and NaHCO. on disestionin the stomach of weaned calves. J. egri. Sci.(Camb.) 9l 497,503.LEIBHOLZ, J., KELLAWAY, R. C. andHARGREAVE, G. T. 1980. Effects of sodiumchloride and sodium bicarbonate in the diet onthe performance in calves. Anim. Feed Sci.Technol. 5: 309-314.MAHADEVAN, S., SAUER, F. D., ERFLE,J. D., TEATHER, R. M. and MORSE, P. M.1982. Changes in ammonia concentration, bac-terial counts, pH and volatile fatty acid concen-tration in rumen of cows fed alfalfa hay or con-centrate: urea-corn silage. Can. J. Anim. Sci.62:249-258.

MERCHEN, N., HANSON, T. and KLOP-FENSTEIN, T. 1919. Ruminal by-pass ofbrewers dried grains protein. J. Anim. Sci. 49:l 92-1 98.NICHOLSON, J. W. G., CUNNINGHAM, H.M. and FRIEND. D. W. 1963a. Effect of add-ing buffers to all concentrate rations on feedlotperfbrmance of steers, ration digestibility andintrarumen environment. J. Anim. Sci.22:368-373.NICHOLSON, J. W. G., CUNNINGHAM, H.M. and FRIEND. D. W. 1963b. The additionof buffers to ruminant rations. III. The effectof additions of sodium bicarbonate, sodium pro-pionate, limestone and cod liver oil on apparentdigestibility and nitrogen retention of an all-concentrate ration. Can. J . Anim. Sci. 42:82-81.OKEKE, G. C., BUCHANAN-SMITH, J, G.and GROVUM. W. L. 1983. Effects of bufferson ruminal rate of passage and degradation ofsoybean meal in steers. J. Anim. Sci.56:1393 1399.PHILLIP. L. E. and BUCHANAN-SMITH. J.G. 1982. Effects of ensiling upon free aminoacids and amines in whole-plant-corn, and onits subsequent nutritive value for lambs. Can.J. Anim. Sci. 62: 259-261 .PHILLIP, L. E., BUCHANAN-SMITH, J. G.and GROVUM, W. L. 1981. Food rntake andruminal osmolality in sheep: differentiation ofthe effect of osmolality from that of the prod-ucts of maize silage fermentation. J. Agric. Sci.(Camb.) 96: 439-445.TRENKLE. A. H. 1979. Sodium bicarbonatePages 1-82 rn Beef nutrition. National Feed In-gredient Assoc. Lit. Review. N.F.I.A., DesMoines, Iowa.VAN SOEST, P. J. 1963. Uses of detergentsin the analysis of fibrous feeds. II. A rapidmethod for the determination of fibre and lig-nin. J. Assoc. Off. Anal. Chem. 50: 50.

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