Use of sodium bicarbonate, offered free choice or blended into the ration, to reduce the risk of ruminal acidosis

in cattle
Laura J. Paton1, Karen A. Beauchemin2, 4, Douglas M. Veira3, and Marina A. G. von Keyserlingk3
1Animal Welfare Program, University of British Columbia, Vancouver, British Columbia, Canada V6T 1Z4; 2Agriculture and Agri-Food Canada, Research Center, Lethbridge, Alberta, Canada T1K 4H3; and 3Agriculture and Agri-Food Canada, Research Center, Kamloops, British Columbia, Canada V2B 8A9.

Received 9 February 2006, accepted 6 June 2006.

Paton, L. J., Beauchemin, K. A., Veira, D. M. and von Keyserlingk, M. A. G. 2006. Use of sodium bicarbonate, offered free choice or blended into the ration, to reduce the risk of ruminal acidosis in cattle. Can. J. Anim. Sci. 86: 429437. A study was conducted to determine whether feeding sodium bicarbonate (SB) reduces the risk of subacute ruminal acidosis in cattle fed high concentrate feedlot finishing diets. The experiment was conducted as a replicated 3 3 Latin square design with two squares and 2-wk periods. Three mature, non-lactating Holstein cows were allocated to square 1 and three mature Jersey steers were allocated to square 2. The cattle were ruminally cannulated and gradually adapted to a high concentrate diet before starting the experiment. The basal diet contained approximately 80% stream-rolled barley, on a dry matter (DM) basis, and was offered for ad libitum intake. Treatments were: control (no SB), control diet with cattle given free choice access to a SB mixture consisting of 70% SB and 30% dried molasses (free choice SB), and control diet supplemented with SB (7 g SB kg1 DM; mixed SB). Ruminal pH was measured at the end of each 14-d period for 3 continuous days using an indwelling pH system. Dry matter intake (DMI) was not affected by treatment. However, SB intake depended upon type of cattle and method of provision (P = 0.04); cows had higher SB intake when it was mixed into the diet (57.8 vs. 17.4 g d1), whereas steers had higher SB intake when SB was provided free choice (129.1 vs. 56.1 g d1). Ruminal pH characteristics (mean, maximum, minimum, hours, and area under a threshold pH of 5.8 or 5.5) were not affected by treatment. Although neither method of delivering SB reduced the total time each day that pH was below the pH thresholds used to indicate subacute ruminal acidosis, the number of long (> 4 h) continuous bouts of acidosis (pH 5.8) was reduced (P = 0.01) when SB was mixed into the ration compared with the control. When offered free choice, intake of SB was highly variable among animals and from day-to-day and was inversely correlated to DMI. However, there was no correlation between SB intake and ruminal pH indicating that, when given the opportunity, cattle did not select SB to help prevent ruminal acidosis. Providing SB, either free choice or mixed into the ration, did not eliminate subacute ruminal acidosis in cattle fed high-grain diets. However, mixing SB into the ration reduced the number of long bouts of ruminal acidosis, which could potentially reduce the negative consequences of ruminal acidosis on feed digestion. Key words: Acidosis, beef cattle, high-grain diets, ruminal pH, sodium bicarbonate Paton, L. J., Beauchemin, K. A., Veira, D. M. et von Keyserlingk, M. A. G. 2006. Rduction du risque dacidose du rumen chez les bovins par le bicarbonate de sodium offert satit ou mlang la ration. Can. J. Anim. Sci. 86: 429437. Les auteurs ont entrepris une tude pour savoir si le fait de donner du bicarbonate de sodium (BS) aux bovins en parc dengraissement recevant une ration riche en concentrs rduirait le risque dacidose subaige du rumen. Lexprience sest droule en double carr latin de 3 x 3 comprenant deux enclos et deux priodes de deux semaines. Trois vaches Holstein matures non en lactation ont t places dans le premier enclos tandis que le second accueillait trois bouvillons Jersey matures. Les animaux taient canuls au rumen et on les avait graduellement adapts une ration riche en concentrs avant que lexprience dbute. La ration de base contenait environ 80 % de flocons dorge obtenus la vapeur, selon le volume de matire sche, et les animaux y avaient accs volont. Les traitements taient les suivants : ration tmoin (pas de BS), ration tmoin et libre accs un mlange compos 70 % de BS et 30 % de mlasse dshydrate (BS satit) et ration tmoin enrichie de BS (7 g de BS par kg de matire sche; mlange BS). Les auteurs ont dos le pH du rumen pendant trois journes conscutives grce un pHmtre install demeure la fin de chaque priode de 14 jours. Le traitement naffecte pas lingestion de la matire sche (IMS). Toutefois, la quantit de BS ingre dpend du type de bovin et de la mthode employe pour procurer le supplment (P = 0,04); les vaches ingrent plus de BS quand celui-ci est mlang la ration (57,8 c. 17,4 g par jour) tandis que les bouvillons en consomment davantage quand il est offert satit (129,1 c. 56,1 g par jour). Le traitement ne modifie pas les caractristiques du pH du rumen (moyenne, maximum, minimum, dure en heures et superficie au pH infrieur 5,8 ou 5,5). Comparativement la ration tmoin, la ration contenant le BS diminue (P = 0,01) le nombre de longs (> 4 heures) pisodes dacidose (pH 5,8), bien quaucune des deux mthodes employes pour fournir le BS ne rduise la dure globale de la priode durant laquelle le pH est infrieur aux seuils tablis pour dterminer

4To whom correspondence should be addressed (e-mail: beauchemink@agr.gc.ca).

Abbreviations: DM, dry matter; DMI, dry matter intake; r = Pearson correlation coefficient; SB, sodium bicarbonate; SD, standard deviation, SEM, standard error of the mean; VFA, volatile fatty acids
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CANADIAN JOURNAL OF ANIMAL SCIENCE une acidose subaige durant la journe. Quand les animaux ont libre accs au supplment, lingestion de BS varie considrablement dune journe lautre et est inversement corrle lIMS. Il nexiste cependant aucune corrlation entre lingestion de BS et le pH du rumen, signe que, quand ils en ont loccasion, les bovins ne mangent pas du BS pour contribuer prvenir lacidose du rumen. En procurant du BS satit ou avec la ration aux animaux, on ne met pas fin lacidose subaige du rumen chez les bovins recevant une ration riche en crales. Mlanger le BS la ration rduit cependant le nombre dpisodes prolongs dacidose, ce qui pourrait attnuer les consquences ngatives de lacidose du rumen sur la digestion des aliments. Mots cls: Acidose, bovins de boucherie, rations riches en crales, pH du rumen, bicarbonate de sodium

Acidosis causes economic losses for the North American feedlot industry due to increased costs associated with treatment of sick animals, reduced animal performance, and mortality (Nocek 1997; Owens et al. 1998). Acidosis results from the rapid production of ruminal volatile fatty acids (VFA) and lactic acid arising from ingestion of readily fermentable carbohydrates (Britton and Stock 1986). The degree of severity of acidosis can be variable; cattle exhibit acute systemic acidosis as an overt illness while subacute ruminal acidosis is subtle and difficult to diagnose unless cattle are ruminally cannulated. Subacute ruminal acidosis is characterized by periods during which ruminal pH drops below optimum for growth of cellulolytic bacteria (Owens et al. 1998). Cellulolytic bacteria cannot grow at pH < 6.0 (Russell and Wilson 1996), thus subacute ruminal acidosis is typically characterized in terms of the duration that ruminal pH drops below a threshold value ranging from 5.5 to 6.0 (i.e., pH 5.5 used by Schwartzkopf-Genswein et al. 2004; pH 5.6 used by Bevans et al. 2005; pH 5.8 used by Ghorbani et al. 2002). While brief periods of pH < 6.0 are unlikely to affect ruminal digestion, extended periods (i.e., several hours) of low ruminal pH are detrimental to fibre digestion and rumen function (Calsamiglia et al. 2002). Depressed fibre digestion can reduce the performance of feedlot finishing cattle fed barley diets because about onefourth of the dietary DM is fibre (measured as neutral detergent fibre). Low ruminal pH can also lead to acute systemic acidosis characterized by excessive acidity of the blood (Owens et al. 1998). Subacute ruminal acidosis is widespread in beef (Brown et al. 2000; Beauchemin et al. 2001) and dairy (Oetzel 2004; Plaizier 2004) cattle operations in North America, because of the use of high-grain diets. When mixed into the ration, dietary buffers such as sodium bicarbonate (SB) can elevate ruminal pH (Ghorbani et al. 1989; Solorzano et al. 1989; Zinn 1991), although not always (Russell et al. 1980; Hart and Polan 1984). Bicarbonate, the primary buffer in saliva and ruminal fluid, is a strong neutralizer of reductions in ruminal pH (Kohn and Dunlap 1998). Thus, providing supplemental buffer in the feed has the potential of increasing the buffering capacity of the ruminal fluid. Buffers may also elevate ruminal pH by increasing the liquid dilution rate from the rumen, and hence, the flow of soluble starch and small particles from the rumen (Russell and Chow 1993). Despite the variability in response to SB in research studies, buffers are routinely added to commercial cattle diets as a precautionary measure in the prevention of acidosis. Furthermore, SB is sometimes offered free choice to cattle based on the assumption that ruminants fed high-grain diets

will readily consume SB to attenuate the effects of ruminal acidosis. This practice is based on the hypothesis that ruminants select a diet that promotes optimal ruminal conditions (Parsons et al. 1994; Cooper at al. 1996). Cooper et al. (1996) observed that when given the choice between a highgrain barley-based diet with or without SB, sheep preferred a ration with SB. It was assumed that the SB was consumed to attenuate the effects of ruminal acidosis, although ruminal pH was not actually measured in that study. The absence of a dose-dependant response to SB in that study raises the question of whether the response was truly a feedback mechanism versus preference based on palatability. The observation that sheep prefer foods and solutions containing SB, compounds that attenuate ruminal acidosis, was subsequently confirmed in other studies (Phy and Provenza 1998a, b; James and Kyriazakis 2002). However, research that substantiates that preference for feeds containing SB is in response to ruminal acidosis is lacking, particularly for cattle. Keunen et al. (2003) provided dairy cattle experiencing ruminal acidosis with free choice access to SB and concluded that SB intake was insufficient to significantly elevate ruminal pH. To our knowledge, there has been no work of this kind with high-grain feedlot diets. The objectives of the present study were to determine: (1) whether adding SB to the diet reduces the incidence and severity of ruminal acidosis experienced by feedlot cattle fed high-grain diets, and (2) whether SB can be offered free choice as an effective means of preventing ruminal acidosis. MATERIALS AND METHODS Experimental Design and Treatments The experiment was conducted as a replicated 3 3 Latin square design, with squares (groups of cattle). Square 1 consisted of three mature Jersey steers and square 2 of three mature non-lactating Holstein cows. The cattle had been ruminally fistulated (10 cm i.d., Bar Diamond, Parma, ID) for use in a previous experiment, and for reasons of animal welfare we decided to use existing research animals rather than subjecting additional animals to the surgery required for cannulation. Although these animals were mature and not representative of growing cattle housed in feedlots, the daily intake of these cattle was similar to the expected intake of growing feedlot cattle (Beauchemin et al. 2001; Bevans et al. 2005). Twenty-eight days prior to commencing the experiment, the animals were adapted to the high-grain control diet using a series of five diets with decreasing forage to concentrate ratio. We then monitored ruminal pH in these cattle to ensure their pH profiles were similar to those typically observed for feedlot cattle (Bevans et al. 2005).

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The experiment consisted of three 2-wk periods. In each period, animals were assigned to one of the following treatments: (1) control (no SB), (2) free choice access to a SB mixture (free choice SB), and (3) SB incorporated into the ration (7 g kg1 DM; mixed SB). To enhance its palatability, free choice SB was offered as a mixture consisting of 70% SB (Arm & Hammer, Church and Dwight Co., Inc., Princeton, NJ) and 30% dried molasses (Coppock et al. 1986). The SB supplementation rate was chosen based on the manufacturers recommended dose range of 0.5 to 0.75% of dry matter intake (DMI; Arm and Hammer 2006). The control diet consisted of steam-rolled barley (812 g kg1 DM), whole crop barley silage (120 g kg1 DM), and supplement (68 g kg1 DM), which was provided (5% orts, DM basis) for ad libitum intake (Table 1). Feeding and Animal Care The ration was prepared daily using a feed mixing wagon and offered once daily at 1400 for ad libitum intake (at least 5% orts). When SB was added to the ration (i.e., mixed SB), this was done based on a proportion (i.e., 0.7% of DM) of the DM offered. The amount of SB consumed was calculated by assuming that the proportion of SB in the ration and orts was the same. Animals in the free choice SB group had continuous access to an 8-L bucket, which was replenished daily with 500 g of the SB mixture. The remaining cattle had access to 500 g d1 of dried molasses to maintain familiarity with the buckets in their feed troughs. To determine free choice SB intake, any bedding or ration that was deposited in the buckets was first removed by sieving the orts through a sieve (2.36-mm standard testing sieve, Fisher Scientific, Pittsburgh, PA). The SB orts were dried in a 60C oven for 48 h to determine the DM content (Association of Official Analytical Chemists 1995) and reported intakes of SB were corrected for the 30% dried molasses in the mixture. A 7-d adaptation period occurred prior to the start of the experiment to allow the animals to adjust to their stalls and feed troughs and become familiar with the SB mixture. Cattle were housed in individual stalls bedded with rubber mats and shavings and were provided ad libitum access to fresh water. The animals were let outside for a 1-h period of exercise daily. All cattle were cared for according to the Canadian Council on Animal Care (1993) guidelines. Ruminal pH and Fermentation Variables Ruminal pH was continuously measured for 72 h (except during calibration) from day 11 to 13 using an indwelling pH monitoring system (Penner et al. 2006). The system uses a microprocessor-based pH controller (model PHCN-37, Omega Engineering, Stamford, CT) connected to a submersible pH electrode (S650-CDHF, Sensorex, Garden Grove, CA) by a 9-m cable (PHEH-65-30-ATC, Omega Engineering, Stamford, CT) suspended above the cows. The cable passes through the ruminal cannula plug and extends approximately 30 cm into the rumen. A perforated shroud around the pH electrode allows material to percolate through but prevents the electrode from contacting the ruminal epithelium. Two 900-g weights attached to the electrode

Table 1. Ingredient and chemical composition (mean SD) of the control diet (DM basis) Item Ingredientz Whole-crop barley silagey Steam-rolled barley Ground barley Limestone Salt Dicalcium phosphate Mineral-vitamin premixx Dried molasses Soy oil Chemical composition (g kg1 DM) Dry matter Organic matter Crude protein Neutral detergent fibre Acid detergent fibre 120 812 43.1 14 5 3 0.5 1.7 0.7 752 954 154 253 71 7.1 9.7 9.3 39.8 17.4 g kg1 DM SD

DM, dry matter; SD, standard deviation. zAll ingredients used were pelleted excluding barley silage and steamrolled barley. yComposition (kg1 DM) was 130 g crude protein, 413 g neutral detergent fibre, and 201 g acid detergent fibre (based on three period samples). xMineral-vitamin premix supplied (kg1 dietary DM): 63 mg of Zn, 27 mg of Mn, 15 mg of Cu, 0.3 mg of Se, 0.19 mg of Co, 0.67 mg of I, 6000 IU of vitamin A, 600 IU of vitamin D, and 43 IU of vitamin E.

shroud maintain positioning within the ventral sac. Ruminal pH was measured every 5 s and averaged over 5-min intervals and recorded by a data logger (model CR10, Campbell Scientific, Logan, UT). The electrodes were removed daily from the rumen 1 h prior to feeding when animals were exercised and calibrated with pH 4.0 and pH 7.0 standards. In order to compare the daily ruminal pH profiles of each animal, descriptive statistics were used, including mean pH, maximum pH, minimum pH, cumulative time pH was below pH 5.8 or pH 5.5, and area the pH recording was below a straight line drawn at pH 5.8 or 5.5. The area was calculated by adding the absolute value of the negative deviations in pH from pH 5.8 or 5.5 for each 5-min interval and then expressed as pH units h. The pH thresholds of 5.8 and 5.5 were used to indicate subacute ruminal acidosis. These values were selected based on the principle that desirable ruminal pH is above 6.0 (Russell and Wilson 1996), and threshold pH values used to indicate subacute ruminal acidosis by others range from 5.5 to 5.8 [ i.e., pH 5.5 used by Schwartzkopf-Genswein et al. (2004); pH 5.6 used by Bevans et al. (2005); pH 5.8 used by Ghorbani et al. (2002)]. The degree to which ruminal pH fluctuated was calculated as pHDIFF and pHVAR, where pHDIFF is the difference between the maximum and minimum ruminal pH for an individual for that day and pHVAR is the variation in ruminal pH (measured as the standard deviation associated with the mean ruminal pH). The pH recordings for each animal were also examined for individual bouts of subacute ruminal acidosis. A bout of subacute ruminal acidosis was said to occur when rumen pH was 5.8 for a continuous duration of more than 15 min. Because mean pH was recorded every 5 min, a 15-min dura-

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tion represented three continuous pH values below the threshold, eliminating any possible anomalies. Long bouts of ruminal acidosis were considered to be those exceeding 4 h in continuous duration, based on the study of Calsamiglia et al. (2002). That study showed that continuously low pH for periods exceeding 4 h negatively affected fibre digestion, whereas 30-min drops in pH were less detrimental to feed digestion. At 0, 3 and 6 h post-feeding on day 13, approximately 1 L of rumen contents from each animal was obtained from four sites within the rumen (reticulum, dorsal and ventral sacs, and the mat) and composited. The contents were then strained through a polyester monofilament fabric (Pecap 71180/59, mesh opening 1180 m, Tetko Inc., Scarborough, ON) and a portion of the filtrate was used to measure osmolality immediately, while another portion was stored frozen for analysis of VFA. Laboratory Analysis Feed samples (silage and ration) were collected weekly and DM content was determined by drying the samples at 60C for 48 h. The weekly DM content of the ration was used in the calculation of DMI whereas the weekly DM content of the silage was used to adjust the proportion of silage in the ration. Samples of orts were also collected and analyzed for DM content (60C for 48 h), for use in the calculation of DMI. Weekly dried samples of feeds were composited by period and ground through a sieve with 1-mm holes (standard model 4, Arthur Thomas Co., Philadelphia, PA). Association of Official Analytical Chemists (1995) methods were used to determine analytical DM (Official Method 930.15), organic matter (Official Method 942.05), and acid detergent fibre (Official Method 973.18, without correction for residual ash). Crude protein (6.25 nitrogen) was determined by flash combustion (Carlo Erba Combustion Analyzer, model NA2100 Protein, CE Instruments, Milan, Italy) and neutral detergent fibre (with sodium sulphite and amylase) was determined according to Van Soest et al. (1991) without correction for residual ash. Five millilitres of ruminal fluid filtrate were combined with 1 mL of 25% (wt/vol) of meta-phosphoric acid and VFA were quantified by gas chromatography (model 5890, Hewlett Parkard, Little Falls, DE) with a capillary column (30 m 0.25 mm i.d., 1 phase thickness, bonded PEG, Supelco Nukol, Sigma-Aldrich Canada, Oakville, ON), and flame ionization detection. The oven temperature was 100C for 1 min, which was then ramped by 20C min1 to 140C, and then by 8C min1 to 200C min1, and held at this temperature for 5 min. The injector temperature was 200C, the detector temperature was 250C, and the carrier gas was helium. Osmolality of ruminal fluid was determined within 2 h of sampling by centrifuging (13 000 g for 30 min at 4C) the filtrate and determining freezing point depression using an osmometer (model 5004 Automatic Osmometer, Precision Systems Inc., Fisher Scientific, Pittsburgh, PA).

Statistical Methods Means for DMI and ruminal pH variables were calculated by day, and then analyzed using the mixed model procedure of SAS (Proc Mixed; SAS Institute, Inc. 1999) with repeated measures. The model included the fixed effects of treatment and group (i.e., cows or steers) and their interactions. Animal and period were considered random effects and time (or day) was considered a repeated measure. Data for ruminal fermentation were analyzed similarly, with sampling time as the repeated measure. The covariance structures used were autoregressive or compound symmetry, except for minimum and maximum ruminal pH, in which case, unstructured was the best fit. Correlations between DMI and mean ruminal pH (days 11 to 13), SB intakes and mean ruminal pH (days 11 to 13), and free choice SB intake and DMI (days 1 to 14) were determined using Pearson correlations (r). When the main effect of treatment was significant (P < 0.05), means were separated using contrast statements and significance was declared at P < 0.05. RESULTS Dry matter intake averaged ( SEM) 8.5 0.97 kg d1 and was similar among treatments (P = 0.79) and group (P = 0.34) (Table 2). Mean DMI varied among days (P = 0.002), but the effect of day was similar (P = 0.79) for all treatments. There was an interaction (P = 0.04) between group and method of SB delivery for SB intake. Cows had higher (P < 0.001) SB intake when it was mixed into the diet (57.8 vs. 17.4 g d1) whereas steers had higher (P < 0.001) SB intake when it was provided free choice (129.1 vs. 56.1 g d1). Because of this interaction, means are presented separately for cows and steers in Table 2. Ruminal pH for cows and steers averaged ( SEM) 5.86 0.08 and was similar (P > 0.75) among treatments (Table 3). Mean ruminal pH tended (5.96 vs. 5.77, P = 0.08) to be higher for steers than cows; however, no treatment by group interaction occurred (P = 0.34). Therefore, pooled means for all cattle are presented in Table 3. Ruminal pH did not vary among days (P = 0.61; Fig. 1), indicating that 1 d of measurement would have been sufficient to characterize treatment effects on ruminal pH. No treatment differences or group by treatment interactions (P > 0.24) were observed for the other ruminal pH variables measured (Table 3). Compared with cows, steers had a higher minimum ruminal pH (5.31 vs. 5.20, P = 0.01), less time below ruminal pH 5.5 (3.3 vs. 8.2 h, P = 0.07) and pH 5.8 (8.3 vs. 13.2 h, P = 0.09) and less area below pH 5.5 (3.4 vs. 9.3 pH h, P = 0.05) and below pH 5.8 (12.0 vs. 25.5 pH h, P = 0.02). Averaged over the 3-d period, the cattle experienced 3.1 bouts of subacute ruminal acidosis each day (Table 4). The duration of these bouts was highly variable, ranging in length from 1 to 22.5 h d1. Cows had more bouts of subacute ruminal acidosis each day than steers (4.0 vs. 2.2; P = 0.047), which is consistent with their lower pH profiles. Cattle fed mixed SB had fewer long bouts (P = 0.01) of subacute ruminal acidosis (i.e., pH 5.8 for more than 4 h continuously) than those fed control. No group by treatment

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Table 2. Intake of dry matter (DMI) and sodium bicarbonate for non-lactating Holstein cows (n = 3) and Jersey steers (n =3) fed a high-grain diet with no sodium bicarbonate (Control), free choice sodium bicarbonate (Free Choice SB) or sodium bicarbonate mixed into the ration (Mixed SB) Item Cows DMI (kg d1) Sodium bicarbonate (g d1) Steers DMI (kg d1) Sodium bicarbonate (g d1) Control 9.2 Free choice SB 9.2 17.4 7.9 129.1 Mixed SB 8.3 57.8 8.0 56.1 SEM 1.3 4.2 1.3 6.5 P 0.69 <0.001 0.69 <0.001

7.7

DMI, dry matter intake; SEM, standard error of the mean.

Table 3. Ruminal pH characteristics in cattle (n = 6) fed a high-grain diet with no sodium bicarbonate (Control), free choice sodium bicarbonate (Free Choice SB) or sodium bicarbonate mixed into the ration (Mixed SB) Item Mean Minimumz Maximumz Time pH < 5.8 (h) Area < pH 5.8 (pH h) Time pH < 5.5 (h) Area < pH 5.5 (pH h) pHDIFFy pHVARx Control 5.81 5.28 6.39 11.50 19.69 5.91 6.99 1.12 0.22 Free choice SB 5.88 5.16 6.58 9.81 8.11 4.99 7.01 1.42 0.20 Mixed SB 5.89 5.34 6.45 10.94 18.36 6.31 5.11 1.10 0.24 SEM 0.08 0.11 0.09 2.04 4.28 1.62 2.07 0.12 0.18 P 0.75 0.53 0.37 0.84 0.95 0.84 0.69 0.14 0.91

SEM, standard error of the mean. zMean lowest and highest pH for each day and each animal. ypHDIFF, Difference between the maximum and minimum ruminal pH for each animal for each day. xpHVAR, Variation in ruminal pH measured as the standard deviation associated with the mean pH for each treatment.

interactions occurred (P > 0.42) for the variables reported in Table 4. Thus, even though the steers consumed large quantities of free choice SB, they did not have significantly fewer (P = 0.47) bouts (2.2) of subacute ruminal acidosis than the control steers (3.0). For both cows and steers, the number of ruminal acidosis bouts was consistently lower when SB was mixed into the diet than when no SB was fed. Daily DMI and mean ruminal pH were not correlated for cattle fed the control diet or mixed SB (P > 0.05). We also tested this relationship using the previous days intake, in case there was a delayed effect, but there was no effect (P > 0.05). However, a negative correlation between DMI and ruminal pH was observed for cattle fed free choice SB (P = 0.02, r = 0.53), indicating that animals with higher DMI had lower mean pH. A similar relationship was observed when intake on the preceding day was tested against ruminal pH. Further analysis revealed that this correlation was predominately due to cows (P = 0.04, r = 0.68) and not steers (P > 0.05). The correlation between SB intake and mean ruminal pH was examined using the data for steers, because free choice SB intakes of cows were so low. Daily intakes of SB and ruminal pH were not correlated for free choice SB or mixed SB (P > 0.05). In fact, daily free choice SB intake was correlated to DMI (P = 0.002, r = 0.40), with higher intake of free choice SB by animals with lower DMI. This inverse relationship between DMI and free choice SB intake was evident for cows (P = 0.08, r = 0.26) and steers (P < 0.001, r = 0.52).

There were no effects of treatment on VFA concentration or proportions (Table 5). Rumen fluid osmolality was similar among treatments (P = 0.41), and steers had higher osmolality than cows (338 vs. 326 mmol L1; P = 0.04). DISCUSSION The high-grain diet used in this study resulted in relatively low ruminal pH profiles, with a mean pH of 5.86 and pH remaining below 5.8 for about 10.5 h d1. Low ruminal pH profiles are typical for cattle fed feedlot finishing diets containing barley (Ghorbani et al. 2002; Bevans et al. 2005). Low ruminal pH was consistent with other signs of ruminal acidosis, including occasional diarrhea (personal observation) and fluctuating feed intake (Owens et al. 1998). Sodium bicarbonate was added to the ration or offered free choice to determine whether the severity of ruminal acidosis in feedlot cattle could be reduced by providing dietary buffers. Dietary SB has the potential to elevate ruminal pH as it is a strong buffer (Kohn and Dunlap 1998). It can also increase the liquid dilution rate from the rumen, and hence, the flow of soluble starch and small grain particles from the rumen (Russell and Chow 1993). A reduction in the quantity of ruminally degraded starch could be beneficial for feedlot cattle consuming diets based on barley grain, because rapid fermentation of barley starch leads to low ruminal pH (Beauchemin et al. 2001). Adding SB to the ration ensured the desired level of SB was consumed, but offering SB free choice resulted in inconsistent SB intakes. Offering SB free choice, rather than

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Fig. 1. Mean ruminal pH profiles of cattle fed control (solid black line), free choice sodium bicarbonate (dotted line) or sodium bicarbonate mixed into the ration (gray line). Mean ruminal pH was similar among treatments (P = 0.75) and there was no day effect (P = 0.61). Animals were fed once daily at 1400.

Table 4. Number of daily bouts of subacute ruminal acidosis in cattle (n = 6) fed a high-grain diet with no sodium bicarbonate (Control), free choice sodium bicarbonate (Free Choice SB) or sodium bicarbonate mixed into the ration (Mixed SB) Bouts of acidosis Total number of bouts d1 Number of long bouts > 4 h d1 Control 3.50 0.78a Free choice SB 3.28 0.56ab Mixed SB 2.56 0.33b SEM 0.53 0.13 P 0.44 0.04

a, b Within row, means without a common letter differ (P < 0.05).

Table 5. Mean ruminal fermentation characteristics for non-lactating Holstein cows and Jersey steers (n = 6) fed a high concentrate diet with no sodium bicarbonate (Control), free choice sodium bicarbonate (Free Choice SB) or sodium bicarbonate mixed into the ration (Mix SB) Item Volatile fatty acids Total (mM) Acetate (A) (%) Propionate (P) (%) A:P Osmolality (mmol L1) Control 16.8 53.1 28.5 1.9 327 Free choice SB 114.1 53.2 30.5 1.8 333 Mix SB 119.3 51.8 28.2 1.9 336 SEM 11.4 2.8 2.4 0.2 4.5 P 0.82 0.92 0.76 0.86 0.41

mixing it into the feed, would be an economical feeding strategy if effective, in that it avoids unnecessary supplementation. This approach of supplying SB free choice assumes that ruminants will select a diet that elevates ruminal pH when it is below optimum for the rumen microorganisms (Cooper et al. 1996; James and Kyriazakis 2002). Thus, the free choice SB treatment was investigated in the present study to determine whether animals would consume SB when offered free choice, and then whether intake of SB was in response to ruminal acidosis. In past studies, it was presumed that variation in free choice SB intake was related to the degree of ruminal aci-

dosis experienced by individual animals (Cooper et al. 1996; Phy and Provenza 1998a), although ruminal pH was not measured in those studies. In the present study, continuous monitoring of ruminal pH revealed that inconsistent SB intake among animals and from day-to-day was not related to ruminal acidosis. For example, ruminal pH was lower in cows than steers, yet steers consumed more SB than cows when it was offered free choice. The steers consumed SB free choice at levels that exceeded recommendations, but the cows consumed far less. The reason for the difference in free choice SB intake between cows and steers is not known, but it could be related to the previous feeding history of

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these animals. The steers had recently been used in a study in which they received feedlot finishing diets, whereas the cows had previously been fed a high forage diet. It is possible that the cows would have increased their free choice SB intakes had they received the feedlot finishing diet over a longer period of time than the short duration of this experiment. Alternatively, the negative relationship between free choice SB intakes and DMI could indicate that the cattle preferred to consume the diet, rather than SB. It is possible that free choice SB intakes would have been higher had feed intake been restricted. Even when free choice intake of SB exceeded the recommended level (i.e., in the case of steers) there was no correlation between SB intake and ruminal pH. The lack of correlation between these variables indicates that the variation in intake of free choice SB was not related to ruminal pH. Free choice intake of SB did not increase in proportion to low ruminal pH, and high intake of SB did not elevate ruminal pH. Keunen et al. (2003) reported results with dairy cows that also refute the hypothesis that animals will consume SB, when offered free choice, to alleviate ruminal acidosis. In their study, dairy cows were subjected to a ruminal acidosis challenge by replacing 25% of the ration with pellets containing a mixture of ground barley and wheat. The cows were then offered free choice SB to determine whether they would consume SB and alleviate the ruminal acidosis challenge. Despite cows exhibiting ruminal acidosis, intakes of free choice SB were low (27 g d1), and ruminal acidosis was not prevented. Thus, providing SB free choice is not recommended because the inconsistent intake of SB among cattle is not related to differences in ruminal pH. Regardless of method of delivery, SB did not elevate mean ruminal pH and had no effect on the total time or area that ruminal pH was below the pH thresholds used to describe ruminal acidosis. The lack of effect of SB in our study is a concern because SB is widely used in commercial beef and dairy cattle operations. Despite widespread use of SB, the effects of feeding SB on ruminal pH are inconsistent in the scientific literature. For example, Hart and Polan (1984) reported for calves fed a corn based diet with added SB (1.5, 3.0, and 4.5% of dietary DM), ruminal pH was not affected; however, pH values remained relatively high in that study. Similarly, Russell et al. (1980) reported ruminal pH of steers fed a finishing diet based on corn and supplemented with 0.9% SB (DM basis) averaged 6.5 and did not differ from control. In contrast, other studies have reported that the addition of buffers to high concentrate diets increased ruminal pH. Ghorbani et al. (1989) reported that adding 1% SB (DM basis) to a lactating dairy cow diet (60% corn grain) significantly increased mean ruminal pH (from 6.06 to 6.24). Zinn (1991) reported feedlot cattle fed a finishing diet of steam-flaked corn supplemented with 0.75% SB (DM basis) had a higher mean ruminal pH (6.23) compared with cattle fed a non-buffered diet (5.87). Sodium bicarbonate had little effect on elevating ruminal pH in the present study, even though subacute ruminal acidosis was prevalent. Russell and Chow (1993) argue that SB has little impact on ruminal pH because the rumen is satu-

rated with CO2. However, work by Kohn and Dunlap (1998) clearly demonstrates that SB can directly neutralize ruminal pH even under high partial pressure of CO2. More recent models predict that ruminal pH is regulated primarily by the difference in the sum of concentrations of soluble cations (mainly Na+ and K+) and anions (mainly Cl and partially dissociated VFA) (Singh 2005). In the present study, there was no effect of SB on total concentration and proportions of VFA, or osmolality of ruminal fluid; therefore, concentrations of partially dissociated VFA were likely similar for all treatments. Lack of treatment effects on ruminal osmolality would indicate that SB did not alter the flow of potentially degradable starch from the rumen. Therefore, no increase in ruminal pH due to decreased ruminal fermentation of starch would have occurred in the present study (Russell and Chow 1993). The lack of response to SB in the present study was likely due to the low SB intake relative to the additional buffering capacity required to elevate ruminal pH. It is possible that SB intake affected ruminal pH during parts of the day, which was detected in terms of bouts of ruminal acidosis. Thus, in addition to mean pH and time and area below pH 5.8 and 5.5, we also examined the incidence of individual bouts of ruminal acidosis occurring within a day. Quantifying the number and length of individual bouts of subacute ruminal acidosis is important because long, continuous periods of low pH are harmful to rumen function and animal health (de Veth and Kolver 2001; Keunen et al. 2002). In the present study, the only beneficial effect of providing SB on alleviating ruminal acidosis was a reduction in the number of long bouts of ruminal acidosis when SB was mixed into the ration. A reduction in the duration of continuous periods of low pH has been shown in vitro to be beneficial to microbial growth and efficiency, and consequently fibre digestion (Russell and Wilson 1996; Calsamiglia et al. 2002). Thus, when comparing the effectiveness of methods of SB supplementation for ruminal acidosis control, mixed SB was more effective than free choice SB in preventing long bouts of ruminal acidosis, although in both cases, the effects of SB supplementation on preventing ruminal acidosis were subtle. Ruminal pH patterns remained consistent from day to day; however, wide ranges in ruminal pH within a day were observed for all animals irrespective of treatment (Fig. 1). Daily fluctuations in ruminal pH are typical of cattle fed high-grain diets (Cooper and Klopfenstein 1996; Hristov et al. 2001). The variation and magnitude of the daily ruminal pH fluctuations observed in the present study (pHVAR and pHDIFF) were comparable to values for cattle fed a highgrain diet (Cooper and Klopfenstein 1996), but providing SB had no effect on minimizing these fluctuations. After feeding, the pH tended to remain constant or increase for a couple of hours, which is in contrast to the pattern typically reported for cattle fed once daily. Ruminal pH profiles usually decrease immediately following feeding reaching a nadir 8 to 16 h post-feeding, then subsequently increase overnight, peaking prior to feeding (Beauchemin et al. 2001; Keunen et al. 2002; Nocek et al. 2002). In this study, the delayed drop in ruminal pH after feeding was

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likely because of the early afternoon feeding time, which is typically characterized by low feeding activity (Gonyou and Stricklin 1984). Other than a delay in post-prandial pH decline, the pH profiles were typical with nadir pH values occurring between 2300 and 0400, approximately 9 to 13 h post-feeding. Ruminal pH gradually increased and peaked at around 0800 and then slowly declined again until the next feeding. Sodium bicarbonate, offered either free choice or mixed into the ration, had no effect on the diurnal pattern of ruminal pH. The inverse relationship between DMI and ruminal pH indicated that animals with high feed intakes were at greater risk for ruminal acidosis. Thus, differences between cows and steers for time and area below ruminal pH 5.8 and 5.5 likely resulted from cows having a higher feed intake. Reductions in feed intake are common when cattle are fed high-grain diets and experience ruminal acidosis. In this study, offering SB free choice or mixed into the diet had no effect on DMI, and thus did not attenuate the negative effects of ruminal acidosis on DMI even in the case of the steers where free choice SB consumption was high. The effects of SB on DMI have been inconsistent in previous studies. While some work has reported no effect of SB on DMI (Hart and Polan 1984; Coppock et al. 1986; Ghorbani et al. 1989), other researchers have reported that dietary buffers improved feed intake and increased animal performance (Fulton et al. 1979b; Solorzano et al. 1989; Zinn 1991). Keunen et al. (2003) reported that offering free choice SB to dairy cows with ruminal acidosis did not increase DMI, similar to our observations. The cyclical or erratic feeding pattern often displayed by cattle exhibiting ruminal acidosis was apparent in this study as indicated by the significant effect of day on DMI. It has been suggested that cattle adjust feed consumption and intake patterns in attempts to attenuate ruminal acidosis and regulate high levels of VFA in the rumen (Slyter 1976; Owens et al. 1998). Fluctuating feed intake patterns have been documented in other studies with ruminants fed highgrain diets (Fulton et al. 1979a; Cooper et al. 1999; Schwartzkopf-Genswein et al. 2003). Providing SB free choice or mixed into the diet did not reduce the daily variation in DMI. CONCLUSIONS Cattle fed a barley-based feedlot finishing diet experienced subacute ruminal acidosis, characterized by a mean pH of 5.86 and pH profiles below the critical threshold of 5.8 for more than 10 h d1. Providing SB, either mixed into the ration or offered free choice, did not elevate mean ruminal pH or reduce the total duration of ruminal acidosis each day. However, mixing SB into the ration was slightly more effective than providing it free choice because it reduced the number of long bouts of ruminal acidosis each day. Long bouts of ruminal acidosis are detrimental to microbial growth and efficiency, and consequently fibre digestion. Intake of free choice SB was highly variable amongst animals and from day to day, but this variability was not related to ruminal pH. Free choice intake of SB did not increase in proportion to low ruminal pH, and high intake of SB did

not elevate ruminal pH. This study refutes the hypothesis that ruminants will consume SB to attenuate the effects of ruminal acidosis; however, longer-term studies need to be conduced before fully rejecting this possibility. ACKNOWLEDGEMENTS We thank the significant contributions of our technician B. Farr for her assistance and Mr. T. Enz for his statistical advice.
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