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Influence of drinking water with quebracho tannin on intake of endophyte-infected tall fescue by cattle
Animal Feed Science and Technology 188 (2014) 13–16
Contents lists available at ScienceDirect
Animal Feed Science and Technology journal homepage: www.elsevier.com/locate/anifeedsci
Influence of drinking water with quebracho tannin on intake of endophyte-infected tall fescue by cattle Tiffanny Lyman Jensen ∗ , Frederick D. Provenza, Juan J. Villalba Department of Wildland Resources, Utah State University, Logan, UT 84322-5230, United States
a r t i c l e
i n f o
Article history: Received 12 February 2013 Received in revised form 8 November 2013 Accepted 13 November 2013
Keywords: Tannin Endophyte-infected tall fescue Cattle Intake
a b s t r a c t Endophyte-infected tall fescue (TF) contains alkaloids that can adversely affect the productivity and health of livestock. These effects are exacerbated when livestock forage on TF grown in monoculture, but they are mitigated when livestock forage on pastures that contain TF in mixture with tannin-containing plants, evidently because tannins bind to the steroidal and protein-like alkaloids found in TF. Cattle and sheep will drink water that contains up to 20 g/L condensed tannins. We hypothesized that cattle (n = 8) offered water containing quebracho tannins (10 g/L) would eat more TF than cattle (n = 8) offered plain water. During the trial, cattle were offered 9 kg/(animal * d) of freshly harvested TF. Cattle drank less water with than without tannins (17.7 kg/d vs. 24.6 kg/d; P < 0.005), and they tended to eat less TF (4.3 kg/d vs. 5.1 kg/d; P = 0.10). No differences in weight change occurred during the trial (P = 0.83) or between groups (P = 0.70). The negative impacts of tannins on water intake likely depressed intake of TF, but the lower intakes of tannin water and TF did not adversely affect BW. On the one hand, water restriction and dehydration reduce food intake and any agent that limits intake of water will adversely affect food intake and animal performance. On the other hand, while cattle with fresh water maintained BW during the trial, so did cattle with tannin water, which suggests the tannin may have benefited the cattle fed a diet of orchard grass hay and TF. Further studies are required to assess the role of tannins in water on both the intake and performance of livestock eating TF. © 2013 Elsevier B.V. All rights reserved.
1. Introduction Land managers are constantly looking for ways to improve production of forages and livestock in ways that maintain ecological integrity and reduce costs of production. Discovering ways to sustainably feed people is important for producers increasingly challenged to meet global needs for food and to maintain the integrity of natural systems. One way to meet both environmental and economic objectives is to use plants such as endophyte-infected tall fescue (TF) that thrive under stressful environmental conditions including drought and heavy grazing. Interestingly, plants such as TF often contain compounds that limit intake by herbivores and may cause health problems when eaten in large amounts. Endophyte-infected TF contains alkaloids that benefit the plant but can adversely affect the health of livestock, especially when TF is grown in monoculture as opposed to mixtures with legumes that enhance its forage value. Plants such as birdsfoot trefoil contain condensed tannins that increase livestock intake of TF and mitigate the negative effects of alkaloids in TF (Lyman et al., 2008, 2011, 2012, 2013;
∗ Corresponding author. Tel.: +1 435 797 3576; fax: +1 435 797 3796. E-mail address: [email protected] (T.L. Jensen). 0377-8401/$ – see front matter © 2013 Elsevier B.V. All rights reserved. http://dx.doi.org/10.1016/j.anifeedsci.2013.11.005
14
T.L. Jensen et al. / Animal Feed Science and Technology 188 (2014) 13–16
Villalba et al., 2011; Owens et al., 2012a,b). Tannins have a high affinity for binding to the steroidal and protein-like alkaloids in TF (Jones and Mangan, 1977; Malinow et al., 1979; Okuda et al., 1982). Cattle and sheep drink water that contains up to 20 g/L condensed tannins (Kronberg, 2008, 2010). These findings suggest livestock may increase their preference for and intake of forages such as TF high in alkaloids if they are provided with water containing tannins, as tannins may mitigate the negative physiological effects of alkaloids. We hypothesized cattle would drink water with tannins and that tannin in water would enable cattle to consume more TF than animals given only water. 2. Materials and methods 2.1. Adaptation and trial The study took place at the Poisonous Plants Research Facility in Richmond, UT, USA (41 55 N 111 48 W). Throughout the 19-d experiment, 16 fall-born calves (248 kg ± 44 kg initial average body weight [BW]) were housed indoors in individual, adjacent pens measuring 4.5 m × 6.09 m. During the 10-d adaptation period, when calves were given time to adjust to their pens and the feeding and watering regimes, they were fed orchard grass hay and offered water with increasing amounts of tannin: 2.5 g/L (3 d), 5 g/L (2 d), 7.5 g/L (2 d), and 10 g/L (3 d) tannins. During the ensuing 9-d trial, cattle in one group (n = 8) had 25 L/d of water containing quebracho tannins (10 g/L), while cattle in the other group (n = 8) had 25 L/d of water without tannins. Tannin concentrations were based on the work of Kronberg (2008), who found that cattle and sheep readily drink water containing tannins at concentrations up to 20 g/L of their daily food intake. Water was offered in 95 L buckets placed opposite each food trough. Cattle in both groups were fed 9 kg/animal of freshly harvested TF from 0700 to 0830. Foods were offered in individual 68 L containers secured within each pen. Tall fescue and water refusals were weighed between 0830 and 0930. Following each morning trial, each calf was fed 5 kg of orchard grass hay, which all calves ate in entirety. Cattle were weighed, after an overnight fast, prior to and after the 19-d trial. All procedures followed the protocols approved by animal care and use (IACUC protocol approval number 1372). 2.2. Tall fescue In 2006, monocultures of tall fescue (TF) (Festuca arundinaceae, Kentucky 31 endophyte-infected) were planted at the Utah Agric. Exp. Stn. Pasture Research Facility in Lewiston, UT (41 57 N. 111 52 W.). The TF was harvested (20–30 cm high and in a vegetative state) each morning at 0600 and transported it to the animal research facility. Representative samples of TF were hand-harvested at 25–50 mm above ground level at the end of the study, placed in plastic bags covered with dry ice, and transported to a freezer where they were kept at −20 ◦ C. They were subsequently freeze dried, ground through a Wiley mill with a 1-mm screen, and analyzed for dry matter (Method 930.15; AOAC, 2000) neutral detergent fiber (NDF) (Van Soest et al., 1991), nitrogen (N) (Method 990.03; AOAC, 2000), and ergovaline (using a high pressure liquid chromatography [HPLC] method; Rottinghaus et al., 1991), one of the primary alkaloids in TF that causes toxicosis in cattle (Yate and Powell, 1988). 2.3. Statistical analyses Food and water intake and BW were analyzed using a mixed-effects model that accounted for the random effect of animal within group, and the fixed effects of group and day and their interaction. The variance–covariance structure used was the one which yielded the lowest Bayesian information criterion. Day entered the model as a repeated measure. Analyses were computed using the MIXED procedure of SAS (SAS Institute Inc., Cary, NC; Version 9.1 for Windows). The model diagnostics included testing for a normal distribution of the error residuals and homogeneity of variance. 3. Results Cattle given quebracho tannin in their water drank less water than cattle that drank only fresh water (17.7 kg/d vs. 24.6 kg/d; P < 0.005; Fig. 1). They also tended to eat less TF than cattle offered fresh water (4.3 kg/d vs. 5.1 kg/d; P = 0.10; Fig. 2). Intake of TF varied across days (P < 0.0001), but Group did not interact with Day (P = 0.43; Fig. 1). Tall fescue contained 559 g/kg (±24) NDF, 113 g/kg (±13) CP, and 247 g/kg (±24) ergovaline. Orchard grass hay contained 560 g/kg (±5) NDF, and 75 g/kg (±19) CP. Cattle given fresh water had an initial average BW of 253 kg (±44 kg) and they maintained that weight to the end of the trial (253 ± 39 kg; P = 0.83). Cattle given tannin water had an initial average BW of 244 kg (±47 kg) and an average BW of 252 kg (±42 kg) post trial (P = 0.83). No differences in cattle weights were observed between groups during the trial (P = 0.70). Cattle of 250 kg BW require 20.3 MJ/d of NEm (Net energy of maintenance) and 239 g/d of metabolizable protein (MP) to cover their maintenance requirements (NRC, 2000). Cattle ate between 9 and 10 kg of DM (4–5 kg TF and 5 kg of orchard grass) which provided enough NE (26.2 MJ/d) and MP (660 g/d) to cover requirements for maintenance and growth (NRC, 2000).
T.L. Jensen et al. / Animal Feed Science and Technology 188 (2014) 13–16
15
Fig. 1. Intake of fresh water or water containing 10 g/L quebracho tannins by cattle. Bars are standard errors.
Fig. 2. Intake of fresh cut endophyte-infected tall fescue by cattle offered fresh water or fresh water containing 1% quebracho tannins. Bars are standard errors.
4. Discussion In three studies, Kronberg (2008, 2010) found that cattle and sheep would drink water that contains quebracho tannins at concentrations up to 20 g/L of their daily water intake. Based on these studies, we hypothesized cattle with or without quebracho tannin in water would drink similar amounts of water, but cattle with tannin in their water would eat more TF compared with cattle given plain water because tannins bind to alkaloids which reduces the negative postingestive effects of alkaloids. Contrary to our hypothesis, cattle with tannins in water drank less water than cattle with fresh water, and they also tended to eat less TF. We suggest two explanations for these findings. On the one hand, dry matter intake decreases as water intake decreases (MacFarlane and Howard, 1972; Silanikove, 1987). Water restriction and dehydration reduce food intake and any agent that limits intake of water will adversely affect food intake and animal performance (Balch et al., 1953; Silanikove, 1985). On the other hand, while cattle with fresh water maintained BW during the trial (253 ± 44 kg vs. 253 ± 39 kg), so did cattle with tannin water (244 kg ± 47 kg vs. 252 ± 42 kg), which suggests the tannin may have benefited the cattle fed a diet of orchard grass hay and TF. Tannins can improve protein utilization in ruminants (Owens et al., 2012a, b), which can enhance detoxification processes (Foley and McArthur, 1994; Foley et al., 1999) by providing limiting amino acids that increase metabolism and excretion of secondary compounds like the alkaloids in TF (Illius and Jessop, 1995). In addition, by increasing the supply of protein to the small intestine, tannins may have improved the balance of protein to energy and reduced the need to eat more TF in order to meet needs for limiting nutrients in the diet (Provenza and Villalba, 2006). Nevertheless, tannins at high doses can negatively impact the energy and protein metabolism of the herbivore (Illius and Jessop, 1995), and tannins in water evidently limited intake of water, which likely limited intake of TF.
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T.L. Jensen et al. / Animal Feed Science and Technology 188 (2014) 13–16
Tannins can increase mean retention times of both solutes and particles in the digestive tract (Hofmann, 1989; Silanikove et al., 2001), thus increasing the volume and weight of digested materials even when water and feed intake are decreased. Thus, cattle ingesting water with tannins may have been retaining digested food, thus showing little or no change in BW. The briefness of the trial and the lack of data on empty-carcass BW limit conclusions regarding BW changes. In light of these reservations, we contend that any beneficial effects of tannins in drinking water for intake of TF and performance of cattle remain to be established. Further studies are required to assess the role of tannins in water on both the intake and performance of livestock. If means can be devised to enable cattle and sheep to meet daily water needs while drinking water with tannins, then further efforts could be made to discover the viability of tannin water for influencing TF consumption. Acknowledgments This research was supported by grants from the Utah Agricultural Experiment Station and the Initiative for the Future of Agriculture and Food Systems, USDA (agreement no. 2001-52103-11215). This paper is published with the approval of the Director, Utah Agricultural Experiment Station, and Utah State University, as journal paper number 8502. References AOAC, 2000. Official Methods of Analysis, 17th ed. Association of Official Analytical Chemists, Gaithersburg, MD. Balch, C.C., Balch, D.H., Johnson, V.W., Turner, J., 1953. Factors affecting the utilization of food by dairy cows: 7. The effect of limited water intake on the digestibility and the rate of passage of hay. Br. J. Nutr. 7, 212–224. Foley, W.J., McArthur, C., 1994. The effects and costs of allelochemicals for mammalian herbivores: an ecological perspective. In: Chivers, D.J., Langer, P. (Eds.), The Digestive System in Mammals: Food, Form and Function. Cambridge University Press, Cambridge, pp. 370–391. Foley, W.J., Iason, G.R., McArthur, C., 1999. Role of plant secondary metabolites in the nutritional ecology of mammalian herbivores: how far have we come in 25 years? In: Jung, H.G., Fahey Jr., G.C. (Eds.), Nutritional Ecology of Herbivores, Proceedings of the Fifth International Symposium on the Nutrition of Herbivores. American Society of Animal Science, Savoy, IL, pp. 130–209. Hofmann, R.R., 1989. Evolutionary steps of ecophysiological adaptation and diversification of ruminants: a comparative view of their digestive system. Oecologia 78, 443–457. Illius, A.W., Jessop, N.S., 1995. Modeling metabolic costs of allelochemical ingestion by foraging herbivores. J. Chem. Ecol. 21, 693–719. Jones, W.T., Mangan, J.L., 1977. Complexes of condensed tannins of sainfoin (Onobrychis viciifolia Scop.) with fraction-1 leaf protein and with submaxillary mucoprotein, and their reversal by polyethylene-glycol and pH. J. Sci. Food Agric. 28, 126–136. Kronberg, S.L., 2008. Intake of water containing condensed tannin by cattle and sheep. Rangeland Ecol. Manage. 61, 354–358. Kronberg, S.L., 2010. Sheep ingestion of water containing quebracho and black wattle tannin. Rangeland Ecol. Manage. 63, 258–262. Lyman, T.D., Provenza, F.D., Villalba, J.J., 2008. Sheep foraging behavior in response to interactions among alkaloids, tannins, and saponins. J. Sci. Food Agric. 88, 824–831. Lyman, T.D., Provenza, F.D., Villalba, J.J., Wiedmeier, R.D., 2011. Cattle preferences differ when endophyte-infected tall fescue, birdsfoot trefoil, and alfalfa are grazed in difference sequences. J. Anim. Sci. 89, 1131–1137. Lyman, T.D., Provenza, F.D., Villalba, J.J., Wiedmeier, R.D., 2012. Phytochemical complementarities among endophyte-infected tall fescue, reed canarygrass, birdsfoot trefoil, and alfalfa affect cattle foraging. Animal 6, 676–682. Lyman, T.D., Provenza, F.D., Villalba, J.J., 2013. Influence of diet sequence on intake of foods containing ergotamine d-tartrate, tannins and saponins by sheep. Appl. Anim. Behav. Sci. 144, 57–62. MacFarlane, W.V., Howard, B., 1972. Comparative water and energy economy of wild and domestic mammals. Symp. Zool. Soc. Lond. 31, 251–296. Malinow, M.R., McLaughlin, P., Stafford, C., Livingston, A.L., Kohler, G.O., Cheeke, P.R., 1979. Comparative effects of alfalfa saponins and alfalfa fiber on cholesterol absorption in rats. Am. J. Clin. Nutr. 32, 1810–1812. NRC, 2000. Nutrient Requirements of Beef Cattle, Updated 7th rev. ed. National Academies Press, Washington, DC. Okuda, T., Morie, K., Shiota, M., 1982. Effects of interaction of tannins and coexisting substances: III. Formation and solubilization of precipitates with alkaloids. J. Pharm. Soc. Jpn. 102, 854–858. Owens, J., Provenza, F.D., Wiedmeier, R.D., Villalba, J.J., 2012a. Supplementing endophyte-infected tall fescue or reed canarygrass with alfalfa or birdsfoot trefoil increases forage intake and digestibility by sheep. J. Sci. Food Agric. 92, 987–992. Owens, J., Provenza, F.D., Wiedmeier, R.D., Villalba, J.J., 2012b. Influence of saponins and tannins on intake and nutrient digestion of alkaloid-containing foods. J. Sci. Food Agric. 92, 2373–2378. Provenza, F.D., Villalba, J.J., 2006. Foraging in domestic vertebrates: linking the internal and external milieu. In: Bels, V.L. (Ed.), Feeding in Domestic Vertebrates: From Structure to Function. CABI Publishing, Oxfordshire, UK, pp. 210–240. Rottinghaus, G.E., Garner, G.B., Creighton, N.C., Ellis, J.L., 1991. HPLC method for quantitating ergovaline in endophyte-infested tall fescue: seasonal variation of ergovaline levels in stems with leaf sheaths, leaf blades, and seed heads. J. Agric. Food Chem. 39, 112–115. Silanikove, N., 1985. Effect of dehydration on feed intake and dry matter digestibility in desert (black Bedouin) and non-desert (Swiss Saanen) goats fed on lucerne hay. Comp. Biochem. Physiol. 80A, 449–452. Silanikove, N., 1987. Impact of shade in a hot Mediterranean summer on feed intake, feed utilization and body fluid distribution in sheep. Appetite 9, 207–215. Silanikove, N., Gilboa, N., Nitsan, Z., 2001. Effect of polyethylene glycol on rumen volume and retention time of liquid and particulate matter along the digestive tract in goats fed tannin-rich carob leaves (Ceratonia siliqua). Small Rumin. Res. 40, 95–99. Van Soest, P.J., Robertson, J.B., Lewis, B.A., 1991. Methods for dietary fiber, neutral detergent fiber, and nonstarch polysaccharides in relation to animal nutrition. J. Dairy Sci. 74, 3583–3597. Villalba, J.J., Provenza, F.D., Clemensen, A.K., Larsen, R., Junke, J., 2011. Preference for diverse pastures by sheep in response to intraruminal administrations of tannins, saponins, and alkaloids. Grass Forage Sci. 66, 224–236. Yate, S.G., Powell, R.G., 1988. Analysis of ergopeptine alkaloids in endophyte-infected tall fescue. J. Agric. Food Chem. 36, 337–340.
Contents lists available at ScienceDirect
Animal Feed Science and Technology journal homepage: www.elsevier.com/locate/anifeedsci
Influence of drinking water with quebracho tannin on intake of endophyte-infected tall fescue by cattle Tiffanny Lyman Jensen ∗ , Frederick D. Provenza, Juan J. Villalba Department of Wildland Resources, Utah State University, Logan, UT 84322-5230, United States
a r t i c l e
i n f o
Article history: Received 12 February 2013 Received in revised form 8 November 2013 Accepted 13 November 2013
Keywords: Tannin Endophyte-infected tall fescue Cattle Intake
a b s t r a c t Endophyte-infected tall fescue (TF) contains alkaloids that can adversely affect the productivity and health of livestock. These effects are exacerbated when livestock forage on TF grown in monoculture, but they are mitigated when livestock forage on pastures that contain TF in mixture with tannin-containing plants, evidently because tannins bind to the steroidal and protein-like alkaloids found in TF. Cattle and sheep will drink water that contains up to 20 g/L condensed tannins. We hypothesized that cattle (n = 8) offered water containing quebracho tannins (10 g/L) would eat more TF than cattle (n = 8) offered plain water. During the trial, cattle were offered 9 kg/(animal * d) of freshly harvested TF. Cattle drank less water with than without tannins (17.7 kg/d vs. 24.6 kg/d; P < 0.005), and they tended to eat less TF (4.3 kg/d vs. 5.1 kg/d; P = 0.10). No differences in weight change occurred during the trial (P = 0.83) or between groups (P = 0.70). The negative impacts of tannins on water intake likely depressed intake of TF, but the lower intakes of tannin water and TF did not adversely affect BW. On the one hand, water restriction and dehydration reduce food intake and any agent that limits intake of water will adversely affect food intake and animal performance. On the other hand, while cattle with fresh water maintained BW during the trial, so did cattle with tannin water, which suggests the tannin may have benefited the cattle fed a diet of orchard grass hay and TF. Further studies are required to assess the role of tannins in water on both the intake and performance of livestock eating TF. © 2013 Elsevier B.V. All rights reserved.
1. Introduction Land managers are constantly looking for ways to improve production of forages and livestock in ways that maintain ecological integrity and reduce costs of production. Discovering ways to sustainably feed people is important for producers increasingly challenged to meet global needs for food and to maintain the integrity of natural systems. One way to meet both environmental and economic objectives is to use plants such as endophyte-infected tall fescue (TF) that thrive under stressful environmental conditions including drought and heavy grazing. Interestingly, plants such as TF often contain compounds that limit intake by herbivores and may cause health problems when eaten in large amounts. Endophyte-infected TF contains alkaloids that benefit the plant but can adversely affect the health of livestock, especially when TF is grown in monoculture as opposed to mixtures with legumes that enhance its forage value. Plants such as birdsfoot trefoil contain condensed tannins that increase livestock intake of TF and mitigate the negative effects of alkaloids in TF (Lyman et al., 2008, 2011, 2012, 2013;
∗ Corresponding author. Tel.: +1 435 797 3576; fax: +1 435 797 3796. E-mail address: [email protected] (T.L. Jensen). 0377-8401/$ – see front matter © 2013 Elsevier B.V. All rights reserved. http://dx.doi.org/10.1016/j.anifeedsci.2013.11.005
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T.L. Jensen et al. / Animal Feed Science and Technology 188 (2014) 13–16
Villalba et al., 2011; Owens et al., 2012a,b). Tannins have a high affinity for binding to the steroidal and protein-like alkaloids in TF (Jones and Mangan, 1977; Malinow et al., 1979; Okuda et al., 1982). Cattle and sheep drink water that contains up to 20 g/L condensed tannins (Kronberg, 2008, 2010). These findings suggest livestock may increase their preference for and intake of forages such as TF high in alkaloids if they are provided with water containing tannins, as tannins may mitigate the negative physiological effects of alkaloids. We hypothesized cattle would drink water with tannins and that tannin in water would enable cattle to consume more TF than animals given only water. 2. Materials and methods 2.1. Adaptation and trial The study took place at the Poisonous Plants Research Facility in Richmond, UT, USA (41 55 N 111 48 W). Throughout the 19-d experiment, 16 fall-born calves (248 kg ± 44 kg initial average body weight [BW]) were housed indoors in individual, adjacent pens measuring 4.5 m × 6.09 m. During the 10-d adaptation period, when calves were given time to adjust to their pens and the feeding and watering regimes, they were fed orchard grass hay and offered water with increasing amounts of tannin: 2.5 g/L (3 d), 5 g/L (2 d), 7.5 g/L (2 d), and 10 g/L (3 d) tannins. During the ensuing 9-d trial, cattle in one group (n = 8) had 25 L/d of water containing quebracho tannins (10 g/L), while cattle in the other group (n = 8) had 25 L/d of water without tannins. Tannin concentrations were based on the work of Kronberg (2008), who found that cattle and sheep readily drink water containing tannins at concentrations up to 20 g/L of their daily food intake. Water was offered in 95 L buckets placed opposite each food trough. Cattle in both groups were fed 9 kg/animal of freshly harvested TF from 0700 to 0830. Foods were offered in individual 68 L containers secured within each pen. Tall fescue and water refusals were weighed between 0830 and 0930. Following each morning trial, each calf was fed 5 kg of orchard grass hay, which all calves ate in entirety. Cattle were weighed, after an overnight fast, prior to and after the 19-d trial. All procedures followed the protocols approved by animal care and use (IACUC protocol approval number 1372). 2.2. Tall fescue In 2006, monocultures of tall fescue (TF) (Festuca arundinaceae, Kentucky 31 endophyte-infected) were planted at the Utah Agric. Exp. Stn. Pasture Research Facility in Lewiston, UT (41 57 N. 111 52 W.). The TF was harvested (20–30 cm high and in a vegetative state) each morning at 0600 and transported it to the animal research facility. Representative samples of TF were hand-harvested at 25–50 mm above ground level at the end of the study, placed in plastic bags covered with dry ice, and transported to a freezer where they were kept at −20 ◦ C. They were subsequently freeze dried, ground through a Wiley mill with a 1-mm screen, and analyzed for dry matter (Method 930.15; AOAC, 2000) neutral detergent fiber (NDF) (Van Soest et al., 1991), nitrogen (N) (Method 990.03; AOAC, 2000), and ergovaline (using a high pressure liquid chromatography [HPLC] method; Rottinghaus et al., 1991), one of the primary alkaloids in TF that causes toxicosis in cattle (Yate and Powell, 1988). 2.3. Statistical analyses Food and water intake and BW were analyzed using a mixed-effects model that accounted for the random effect of animal within group, and the fixed effects of group and day and their interaction. The variance–covariance structure used was the one which yielded the lowest Bayesian information criterion. Day entered the model as a repeated measure. Analyses were computed using the MIXED procedure of SAS (SAS Institute Inc., Cary, NC; Version 9.1 for Windows). The model diagnostics included testing for a normal distribution of the error residuals and homogeneity of variance. 3. Results Cattle given quebracho tannin in their water drank less water than cattle that drank only fresh water (17.7 kg/d vs. 24.6 kg/d; P < 0.005; Fig. 1). They also tended to eat less TF than cattle offered fresh water (4.3 kg/d vs. 5.1 kg/d; P = 0.10; Fig. 2). Intake of TF varied across days (P < 0.0001), but Group did not interact with Day (P = 0.43; Fig. 1). Tall fescue contained 559 g/kg (±24) NDF, 113 g/kg (±13) CP, and 247 g/kg (±24) ergovaline. Orchard grass hay contained 560 g/kg (±5) NDF, and 75 g/kg (±19) CP. Cattle given fresh water had an initial average BW of 253 kg (±44 kg) and they maintained that weight to the end of the trial (253 ± 39 kg; P = 0.83). Cattle given tannin water had an initial average BW of 244 kg (±47 kg) and an average BW of 252 kg (±42 kg) post trial (P = 0.83). No differences in cattle weights were observed between groups during the trial (P = 0.70). Cattle of 250 kg BW require 20.3 MJ/d of NEm (Net energy of maintenance) and 239 g/d of metabolizable protein (MP) to cover their maintenance requirements (NRC, 2000). Cattle ate between 9 and 10 kg of DM (4–5 kg TF and 5 kg of orchard grass) which provided enough NE (26.2 MJ/d) and MP (660 g/d) to cover requirements for maintenance and growth (NRC, 2000).
T.L. Jensen et al. / Animal Feed Science and Technology 188 (2014) 13–16
15
Fig. 1. Intake of fresh water or water containing 10 g/L quebracho tannins by cattle. Bars are standard errors.
Fig. 2. Intake of fresh cut endophyte-infected tall fescue by cattle offered fresh water or fresh water containing 1% quebracho tannins. Bars are standard errors.
4. Discussion In three studies, Kronberg (2008, 2010) found that cattle and sheep would drink water that contains quebracho tannins at concentrations up to 20 g/L of their daily water intake. Based on these studies, we hypothesized cattle with or without quebracho tannin in water would drink similar amounts of water, but cattle with tannin in their water would eat more TF compared with cattle given plain water because tannins bind to alkaloids which reduces the negative postingestive effects of alkaloids. Contrary to our hypothesis, cattle with tannins in water drank less water than cattle with fresh water, and they also tended to eat less TF. We suggest two explanations for these findings. On the one hand, dry matter intake decreases as water intake decreases (MacFarlane and Howard, 1972; Silanikove, 1987). Water restriction and dehydration reduce food intake and any agent that limits intake of water will adversely affect food intake and animal performance (Balch et al., 1953; Silanikove, 1985). On the other hand, while cattle with fresh water maintained BW during the trial (253 ± 44 kg vs. 253 ± 39 kg), so did cattle with tannin water (244 kg ± 47 kg vs. 252 ± 42 kg), which suggests the tannin may have benefited the cattle fed a diet of orchard grass hay and TF. Tannins can improve protein utilization in ruminants (Owens et al., 2012a, b), which can enhance detoxification processes (Foley and McArthur, 1994; Foley et al., 1999) by providing limiting amino acids that increase metabolism and excretion of secondary compounds like the alkaloids in TF (Illius and Jessop, 1995). In addition, by increasing the supply of protein to the small intestine, tannins may have improved the balance of protein to energy and reduced the need to eat more TF in order to meet needs for limiting nutrients in the diet (Provenza and Villalba, 2006). Nevertheless, tannins at high doses can negatively impact the energy and protein metabolism of the herbivore (Illius and Jessop, 1995), and tannins in water evidently limited intake of water, which likely limited intake of TF.
16
T.L. Jensen et al. / Animal Feed Science and Technology 188 (2014) 13–16
Tannins can increase mean retention times of both solutes and particles in the digestive tract (Hofmann, 1989; Silanikove et al., 2001), thus increasing the volume and weight of digested materials even when water and feed intake are decreased. Thus, cattle ingesting water with tannins may have been retaining digested food, thus showing little or no change in BW. The briefness of the trial and the lack of data on empty-carcass BW limit conclusions regarding BW changes. In light of these reservations, we contend that any beneficial effects of tannins in drinking water for intake of TF and performance of cattle remain to be established. Further studies are required to assess the role of tannins in water on both the intake and performance of livestock. If means can be devised to enable cattle and sheep to meet daily water needs while drinking water with tannins, then further efforts could be made to discover the viability of tannin water for influencing TF consumption. Acknowledgments This research was supported by grants from the Utah Agricultural Experiment Station and the Initiative for the Future of Agriculture and Food Systems, USDA (agreement no. 2001-52103-11215). This paper is published with the approval of the Director, Utah Agricultural Experiment Station, and Utah State University, as journal paper number 8502. References AOAC, 2000. Official Methods of Analysis, 17th ed. Association of Official Analytical Chemists, Gaithersburg, MD. Balch, C.C., Balch, D.H., Johnson, V.W., Turner, J., 1953. Factors affecting the utilization of food by dairy cows: 7. The effect of limited water intake on the digestibility and the rate of passage of hay. Br. J. Nutr. 7, 212–224. Foley, W.J., McArthur, C., 1994. The effects and costs of allelochemicals for mammalian herbivores: an ecological perspective. In: Chivers, D.J., Langer, P. (Eds.), The Digestive System in Mammals: Food, Form and Function. Cambridge University Press, Cambridge, pp. 370–391. Foley, W.J., Iason, G.R., McArthur, C., 1999. Role of plant secondary metabolites in the nutritional ecology of mammalian herbivores: how far have we come in 25 years? In: Jung, H.G., Fahey Jr., G.C. (Eds.), Nutritional Ecology of Herbivores, Proceedings of the Fifth International Symposium on the Nutrition of Herbivores. American Society of Animal Science, Savoy, IL, pp. 130–209. Hofmann, R.R., 1989. Evolutionary steps of ecophysiological adaptation and diversification of ruminants: a comparative view of their digestive system. Oecologia 78, 443–457. Illius, A.W., Jessop, N.S., 1995. Modeling metabolic costs of allelochemical ingestion by foraging herbivores. J. Chem. Ecol. 21, 693–719. Jones, W.T., Mangan, J.L., 1977. Complexes of condensed tannins of sainfoin (Onobrychis viciifolia Scop.) with fraction-1 leaf protein and with submaxillary mucoprotein, and their reversal by polyethylene-glycol and pH. J. Sci. Food Agric. 28, 126–136. Kronberg, S.L., 2008. Intake of water containing condensed tannin by cattle and sheep. Rangeland Ecol. Manage. 61, 354–358. Kronberg, S.L., 2010. Sheep ingestion of water containing quebracho and black wattle tannin. Rangeland Ecol. Manage. 63, 258–262. Lyman, T.D., Provenza, F.D., Villalba, J.J., 2008. Sheep foraging behavior in response to interactions among alkaloids, tannins, and saponins. J. Sci. Food Agric. 88, 824–831. Lyman, T.D., Provenza, F.D., Villalba, J.J., Wiedmeier, R.D., 2011. Cattle preferences differ when endophyte-infected tall fescue, birdsfoot trefoil, and alfalfa are grazed in difference sequences. J. Anim. Sci. 89, 1131–1137. Lyman, T.D., Provenza, F.D., Villalba, J.J., Wiedmeier, R.D., 2012. Phytochemical complementarities among endophyte-infected tall fescue, reed canarygrass, birdsfoot trefoil, and alfalfa affect cattle foraging. Animal 6, 676–682. Lyman, T.D., Provenza, F.D., Villalba, J.J., 2013. Influence of diet sequence on intake of foods containing ergotamine d-tartrate, tannins and saponins by sheep. Appl. Anim. Behav. Sci. 144, 57–62. MacFarlane, W.V., Howard, B., 1972. Comparative water and energy economy of wild and domestic mammals. Symp. Zool. Soc. Lond. 31, 251–296. Malinow, M.R., McLaughlin, P., Stafford, C., Livingston, A.L., Kohler, G.O., Cheeke, P.R., 1979. Comparative effects of alfalfa saponins and alfalfa fiber on cholesterol absorption in rats. Am. J. Clin. Nutr. 32, 1810–1812. NRC, 2000. Nutrient Requirements of Beef Cattle, Updated 7th rev. ed. National Academies Press, Washington, DC. Okuda, T., Morie, K., Shiota, M., 1982. Effects of interaction of tannins and coexisting substances: III. Formation and solubilization of precipitates with alkaloids. J. Pharm. Soc. Jpn. 102, 854–858. Owens, J., Provenza, F.D., Wiedmeier, R.D., Villalba, J.J., 2012a. Supplementing endophyte-infected tall fescue or reed canarygrass with alfalfa or birdsfoot trefoil increases forage intake and digestibility by sheep. J. Sci. Food Agric. 92, 987–992. Owens, J., Provenza, F.D., Wiedmeier, R.D., Villalba, J.J., 2012b. Influence of saponins and tannins on intake and nutrient digestion of alkaloid-containing foods. J. Sci. Food Agric. 92, 2373–2378. Provenza, F.D., Villalba, J.J., 2006. Foraging in domestic vertebrates: linking the internal and external milieu. In: Bels, V.L. (Ed.), Feeding in Domestic Vertebrates: From Structure to Function. CABI Publishing, Oxfordshire, UK, pp. 210–240. Rottinghaus, G.E., Garner, G.B., Creighton, N.C., Ellis, J.L., 1991. HPLC method for quantitating ergovaline in endophyte-infested tall fescue: seasonal variation of ergovaline levels in stems with leaf sheaths, leaf blades, and seed heads. J. Agric. Food Chem. 39, 112–115. Silanikove, N., 1985. Effect of dehydration on feed intake and dry matter digestibility in desert (black Bedouin) and non-desert (Swiss Saanen) goats fed on lucerne hay. Comp. Biochem. Physiol. 80A, 449–452. Silanikove, N., 1987. Impact of shade in a hot Mediterranean summer on feed intake, feed utilization and body fluid distribution in sheep. Appetite 9, 207–215. Silanikove, N., Gilboa, N., Nitsan, Z., 2001. Effect of polyethylene glycol on rumen volume and retention time of liquid and particulate matter along the digestive tract in goats fed tannin-rich carob leaves (Ceratonia siliqua). Small Rumin. Res. 40, 95–99. Van Soest, P.J., Robertson, J.B., Lewis, B.A., 1991. Methods for dietary fiber, neutral detergent fiber, and nonstarch polysaccharides in relation to animal nutrition. J. Dairy Sci. 74, 3583–3597. Villalba, J.J., Provenza, F.D., Clemensen, A.K., Larsen, R., Junke, J., 2011. Preference for diverse pastures by sheep in response to intraruminal administrations of tannins, saponins, and alkaloids. Grass Forage Sci. 66, 224–236. Yate, S.G., Powell, R.G., 1988. Analysis of ergopeptine alkaloids in endophyte-infected tall fescue. J. Agric. Food Chem. 36, 337–340.