Livestock Science 122 (2009) 345–348
Contents lists available at ScienceDirect
Livestock Science j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / l i v s c i
Short communication
Free-choice mineral consumption in Iberian red deer (Cervus elaphus hispanicus) response to diet deficiencies Francisco Ceacero a,b,c,⁎, Tomás Landete-Castillejos a,b,c, Andrés J. García a,b,c, José A. Estévez a,b,c, Alberto Martinez d, Antonio Calatayud d, Enrique Gaspar-López a,b,c, Laureano Gallego a,b a b c d
Instituto de Investigación en Recursos Cinegéticos, IREC (CSIC, UCLM, JCCM), Campus Universitario s/n, 02071 — Albacete, Spain Departamento de Ciencia y Tecnología Agroforestal y Genética, ETSIA (UCLM), Campus Universitario s/n, 02071 — Albacete, Spain Sección de Recursos Cinegéticos, Instituto de Desarrollo Regional, IDR (UCLM), Campus Universitario s/n, 02071 — Albacete, Spain Laboratorio de Ciencia e Ingeniería de Materiales (CIMA), IDR (UCLM), Campus Universitario s/n, 02071 — Albacete, Spain
a r t i c l e
i n f o
Article history: Received 11 April 2008 Received in revised form 4 August 2008 Accepted 4 August 2008 Keywords: Cervus elaphus hispanicus Foraging theory Free-choice Mineral deficiencies Mineral supplements Nutritional wisdom
a b s t r a c t All mammals have the ability to taste salt, so several studies have focus on mineral selection in the diet. Sodium and phosphorus are usually the most limiting for ungulates and thus some ruminants may regulate their consumption according to mineral needs and content of food sources. The aim of this study was to assess if the amount of minerals consumed is related to daily requirements. It was performed during lactation, examining consumption of 23 adult hinds and their calves. Animals were captive in 10,000 m2 enclosures without pasture. Eleven highly bioavailable mineral compounds were offered in a cafeteria test and weighed weekly. Consumption behaviour was recorded by two cameras from dawn till dusk during the whole lactation period. Incidentally, meal offered had enough minerals to meet daily requirements of all elements except Na and Co. Mineral consumption of supplements was scarce for most elements (less than 10% of daily requirements) except for Na (44%) and Co (36%). Cobaltsupplement was consumed by calves much more than by hinds (90% vs. 10%), probably because requirements for growing ruminants are about 10 times higher than for adults. Animals consumed high percentages of Na and Co but not other minerals, even though flavours of NaCl, KCl and salt-mixed trace minerals are very similar. Thus, consumption seems to reflect physiological needs. © 2008 Elsevier B.V. All rights reserved.
1. Introduction Animals require minerals for their normal life processes, but only some of them are essential minerals: those necessary to support an adequate growth, reproduction, and health throughout the life cycle. Because they cannot be synthesized, minerals are necessarily obtained from diet, and thus animals require a mineral intake for a long-term maintenance of body mineral reserves (McDowell, 2003). All food sources have minerals, although in the wild these are usually present in deficient quantities. Sodium and phosphorus are the most ⁎ Corresponding author. Sección de Recursos Cinegéticos, Instituto de Desarrollo Regional, IDR (UCLM). Campus Universitario s/n, 02071 — Albacete, Spain. Tel.: +34 967 599200x2655; fax: +34 967 599233. E-mail address:
[email protected] (F. Ceacero). 1871-1413/$ – see front matter © 2008 Elsevier B.V. All rights reserved. doi:10.1016/j.livsci.2008.08.002
limiting minerals for ungulates, as they are scarce in plants from most kinds of soils worldwide (McDowell, 2003), but many other trace-elements are also often limiting (Arthington, 2002; Grace and Wilson, 2002; Ward and Lardy, 2005). Optimal foraging theory predicts that animals should be able to determine diet composition and should use foraging tactics that optimise energy intake (Stephens and Krebs, 1986), although as a result of requirements of essential nutrients they should also seek to ingest threshold levels of minerals and other essential nutrients. In fact, all mammals have the ability to taste salt (Denton, 1967), and since Belovsky (1978) first explained diet selection of an ungulate by sodium balance, many studies have focused their attention on the role of mineral nutrition in diet selection by vertebrate herbivores. Thus several studies have shown that animals have evolved the ability to detect contents of some minerals and modify their behaviour (Weeks
346
F. Ceacero et al. / Livestock Science 122 (2009) 345–348
and Kirkpatrick, 1976; McNaughton, 1988, 1990) or regulate their consumption according to their needs (Chládek and Zapletal, 2007; Villalba et al., 2008), at least regarding to sodium and phosphorus. This process was defined as nutritional wisdom (Richter, 1943) and has been criticised by some authors when applied to minerals other than sodium and phosphorus (Muller et al., 1977; McDowell, 1985, 1996). However, other authors have proposed Mg, Zn, Fe or Ca as minerals that ungulates may seek in the diet (Barrows, 1977; Furness, 1988; McNaughton, 1988, 1990; Seagle and McNaughton, 1992; Tracy and McNaughton, 1995; Atwood and Weeks, 2002). Unfortunately, most papers on mineral consumption by ungulates have focus on soil licks and forage composition, which do not allow in most cases to assess discrimination ability separately for each mineral. A particularly interesting stage to study mineral requirements is lactation, as this is a highly demanding stage of minerals for hinds because of milk production. In addition, calves also need great amounts of minerals as lactation influences future body and skeleton size (Gomez et al., 2006), as well as antler size (Gaspar-López et al., 2008) or reproduction (Carrión et al., 2008). That is especially important for males as antler growing requires a strong temporarily skeleton demineralization (Muir et al., 1987). Our own previous studies on a wild ruminant ungulate (Cervus elaphus hispanicus) under outdoor captivity show that both hinds and calves can discriminate among minerals, even thought some of them are mixed with such a strong flavoured excipient as NaCl (Ceacero et al., unpublished data). Moreover, differences in behavioural indices of consumption (frequency of visits and intake time) are not only found towards different minerals, but also between mothers and calves. The aim of this study was to assess if the amount of minerals consumed is related to daily requirements, i.e. if animals consume particularly those minerals which are deficient in the diet. 2. Material and methods This study was carried out at the Experimental Farm of Universidad de Castilla-La Mancha in Albacete. The experiment was performed during 2006 lactation period with 23 adult red deer hinds between 2 and 12 years old and their own calves. Lactation lasted from middle of May to middle of September. Study animals were kept captive in 10,000 m2 enclosures without pasture. Both during gestation and throughout
lactation hinds were maintained on diets based on suggestions by Brelurut et al. (1990), using barley straw and hay from barley, alfalfa, oat, and concentrate (16% protein). Deer had ad libitum access to food and water. Meal consumption was estimated by differences between meal offered and meal retired the following day (i.e., as there was always a surplus of food not consumed, feeding regime was ad libitum). It was 3.5 kg of dry matter per day and animal, similar to amounts indicated by Brelurut et al. (1990). No attempt to record individual intake was conducted. 2.1. Mineral content of diet Meal samples were weekly collected, dried and grinded, and 0.6 g samples were used to assess mineral content. Powder was dissolved with 3 mL of 63% HNO3, 1.5 mL of 37% HCl, and 5.5 mL of Ultrapure deionized water. A second wet digestion was carried out in the microwave oven (CEM MDS2000) under the conditions of 345 kPa for 30 min. After cooling, the digestion solutions were transferred to volumetric flasks and 15 mL of Ultrapure deionized water was added for analysis. Ash samples were then examined with an atomic absorption spectrophotometer (Perkin-Elmer Plasma 400, Boston, MA). Each datum was the mean of 3 measures recorded at 0.3 second intervals (see Landete-Castillejos et al., 2007 for more details on this procedure). Water supplied was also analyzed (data not shown because of its negligible mineral values). 2.2. Mineral supplements A string of containers with different minerals were offered inside a wood shed, safe from wind and rain, in a cafeteriastyle free-choice mineral test. Minerals commonly used as supplements were selected in the inorganic chemical form of highest bioavailability, defined as the portion of the mineral which can be used by the animal to meet its bodily needs (Ward and Lardy, 1995; McDowell, 2005). Minerals were offered in formats and contents recommended for ruminants. Thus, minerals showing a low toxic threshold were mixed with NaCl at concentrations that were a compromise between detectability and safety (Table 1; Ward and Lardy, 2005; Arthington, 2002; McDowell, 2003). Containers were weekly weighted with a Gram Precision AK Eagle 30 (±5 g) portable scale. Total consumption of elements studied (Ca, K, Mg, P, Na, S, Zn, Fe, Co, Cu, and Se) was determined throughout supplement
Table 1 Empirical formulas, mineral concentration and format of minerals offered to red deer in a free-choice cafeteria test. Chemical element
Supplement
Empirical formula
Choice offered
Mineral concentration
Ca Co Cu Fe I K Mg Na P Se Zn
Calcium carbonate Cobaltous sulfate Cupric sulfate Iron carbonate Calcium iodate Potassium chloride Magnesium oxide Sodium chloride Dicalcium phosphate Sodium selenite Zinc sulfate
CaCO3 CoSO4(H2O)7 CuSO4(H2O)5 FeCO3 Ca(IO)3 KCl MgO NaCl Ca2(PO4) Na2SeO3 ZnSO4(H2O)
Pure Mixed Mixed Pure Mixed Pure Pure Pure Pure Mixed Mixed
38.0% 0.4% 1.9% 38.0% 0.8% 40.0% 55.0% 40.0% 18.0% 0.8% 7.2%
on NaCl at 7.5% on NaCl at 7.5% on NaCl at 7.5%
on NaCl at 7.5% on NaCl at 20%
F. Ceacero et al. / Livestock Science 122 (2009) 345–348
347
Table 2 Dietary requirements, mineral composition of meal offered to Iberian red deer during the experiment and consumption of mineral elements from cafeteria test with respect to daily requirements. Nutrient
Reqs a
Meal
% DR b offered
Supplements consumption c
% DR consumed
Ca (%) K (%) Mg (%) P (%) Na (%) S (%) Zn (ppm) Fe (ppm) Co (ppm) Cu (ppm) Se (ppm)
0.50 0.65 0.15 0.30 0.18 0.15 35 50 0.15 15 0.30
0.73 0.99 0.32 0.45 0.05 0.12 36 230 0.03 16 1.30
N 100% N 100% N 100% N 100% 27% N 100% ca. 100% N 100% 20% ca. 100% N 100%
0.007 0.011 0.006 0.003 0.080 0.006 1.1 4.6 0.054 0.894 0.016
1.4% 1.7% 4.0% 1.0% 44.4% 4.0% 3.1% 9.2% 36.0% 6.0% 5.3%
All data is referred to kg of dry matter consumed. Note that Ca, K, Mg, P, Na, and S data is expressed in % while Zn, Fe, Co, Cu, and Se is expressed in ppm. a Daily mineral requirements from Blaxter et al. (1988). b DR = Daily Requirements. c Per day and kilogram of food.
Meal offered to study animals had mineral content adequate to meet red deer daily requirements (based on suggestion by Blaxter et al., 1988) for all elements except Na and Co. Consumption of each mineral element from supplements per kg of dry matter and animal is shown on Table 2. Intake was scarce for almost all elements (less than 10% of daily requirements) except for Na (44%), as expected, and Co (36%). Number of intake events and consumption time showed that NaCl was equally consumed by hinds and calves (51% vs. 49%, respectively). However, CoSO4(H2O)7 consumption was greatly biased towards calves (90% vs. 10%).
simple-stomached animals, with about 3% converted to vitamin B12, although conversion increases when diet is Codeficient (Smith and Marston, 1970). Andrews (1956) showed that lambs are the most sensitive to Co deficiency under grazing conditions, and Marston (1970) showed that requirements for growing lambs are about 10 times the reported values for adults (also recently confirmed for growing beef cattle: Kirchgessner et al., 1998; Stangl et al., 1999; Schwarz et al., 2000). That way, Co free-choice consumption not only response to diet deficiencies, but also hind:calf consumption ratio observed agree with previously published data. Absorption of Co and Fe share a common intestinal mucosal transport system, so they mutually inhibit absorption of the other (McDowell, 2003). Meal offered is greatly rich in Fe because this is an important mineral related to calf growth which is usually deficient in milk (McDowell, 2003; Gallego et al., 2006). Thus, the intake of Fe observed may be also increasing Co demands, which would also explain further the high consumption of Co observed.
4. Discussion
5. Conclusion
Results showed that animals consumed relatively high percentages (related to daily requirements) of Na and Co, precisely the minerals that were deficient in their diet. This suggests that deer are able to assess mineral needs, detect mineral presence or content in the diet and that they modulate intake to meet requirements of only those minerals needed. Sodium is one of the most important minerals in animal physiology (Cunha, 1987). Great amounts of this mineral are needed in diet as most Na ingested is excreted, even if the intestine is able to increase Na absorption during deficiency periods (McDowell, 2003). Considerable quantities of Na may also be lost via secretion in milk (Gallego et al., 2006) and sweat (McDowell, 2003). Ability of mammals to taste salt indicates that Na consumption detected in the experiment responds to expected, especially in high demanding stages such as lactation and early growth. Cobalt is also an essential mineral because of its role in vitamin B12. Ruminal bacteria and algae incorporate Co into vitamin B12, which is used both by microorganisms and animal tissues (McDowell, 2003). Cobalt requirements in ruminants are high as absorption is much less efficient than in
Results clearly show how animals have consumed relatively high percentages of Na and Co, which are the most deficient minerals in meal offered, compared with daily requirements. Flavours of NaCl, KCl and trace minerals offered (Zn, I, Se, and Cu; mixed with NaCl) are quite similar so deer seem to be detecting and specifically seeking these elements, not just consuming based on taste for salt as previously proposed. Beyond sodium, at least cobalt consumption also seems to be controlled by nutritional wisdom.
consumption and atomic weight of each element. Mineral consumption behaviour was recorded along the whole experiment from dawn till dusk by two cameras installed in the shed, as explained in Ceacero et al. (under review) for a study assessing effect of lactation variables on mineral discrimination. 3. Results
Acknowledgements Authors wish to thank Fulgencio Cebrián and Isidoro Cambronero for help in animal care and data collection. This study has been funded by projects PAC-01304298 (JCCM), PET2006-0263 (MCYT), PCI08-0115-8730 (JCCM). F. Ceacero was partly funded by Y. Fierro. References Andrews, E.D., 1956. Cobalt deficiency. N. Z. J. Agr. 92, 239.
348
F. Ceacero et al. / Livestock Science 122 (2009) 345–348
Arthington, J.D., 2002. Essential Trace Minerals for Grazing Cattle in Florida. AN086. EDIS Publication, University of Florida. Atwood, T.C., Weeks, H.P., 2002. Sex- and age-specific patterns of mineral lick use by white-tailed deer (Odocoileus virginianus). Am. Midl. Nat. 148, 289–296. Barrows, G.T., 1977. Research efforts have lagged in free-choice feeding. Anim. Nutr. Health 32, 12–14. Belovsky, G.E., 1978. Diet optimization in a generalist herbivore: the moose. Theor. Popul. Biol. 14, 105–134. Blaxter, K.L., Kay, R.N.B., Sharman, G.A.M., Cunningham, J.M.M., Eadie, J., Hamilton, W.J., 1988. Farming the Red Deer. Department of Agriculture and Fishery of Scotland HMSO, Edimburg. Brelurut, A., Pingard, A., Thériez, M., 1990. Le cerf et son élevage. INRA, Paris. Carrión, D., García, A.J., Gaspar–López, E., Landete-Castillejos, T., Gallego, L., 2008. Development of body condition in hinds of Iberian red deer during gestation and its effects on calf birth weight and milk production. J. Exp. Zool. 309A, 1–10. Chládek, G., Zapl, D., et al., 2007. A free-choice intake blocks in beef cows during the grazing season and in winter. Livest. Sci. 106, 41–46. Cunha, T.J., 1987. Salt and Trace Minerals. Salt Institute, Virginia. Denton, D.A., 1967. Salt Appetite, Handbook of Physiology 1. Am. Physiol. Sot., Washington DC. Furness, R.W.,1988. Predation on ground-nesting seabirds by island populations of red deer Cervus elaphus and sheep Ovis. J. Zool. 216, 565–573. Gallego, L., Landete-Castillejos, T., García, A.J., Sánchez, P.J., 2006. Seasonal and lactational changes in mineral composition of milk from Iberian red deer (Cervus elaphus hispanicus). J. Dairy Sci. 89, 589–595. Gaspar-López, E., García, A.J., Landete-Castillejos, T., Carrión, D., Estévez, J.A., Gallego, L., 2008. Growth of the first antler in Iberian red deer (Cervus elaphus hispanicus). Eur. J. Wildl. Res. 54, 1–5. Gómez, J.A., Landete-Castillejos, T., García, A.J., Sánchez, P.J., Estévez, J.A., Gallego, L., 2006. Effect of lactation on mineral composition of first antler in Iberian red deer (Cervus elaphus hispanicus). Livest. Sci. 105, 27–34. Grace, N.D., Wilson, P.R., 2002. Trace element metabolism, dietary requirements, diagnosis and prevention of deficiencies in deer. N. Z. Vet. J. 50, 252–259. Kirchgessner, M., Schwarz, F.J., Stangl, G.I., 1998. Growth performance of beef cattle fed corn silage-based rations without Cu, Zn, Mn, Co and Se supplementation. J. Anim. Physiol. Anim. Nutr. 78, 141–153. Landete-Castillejos, T., Currey, J.D., Estévez, J.A., Gaspar-López, E., García, A.J., Gallego, L., 2007. Influence of physiological effort of growth and chemical composition on antler bone mechanicals properties. Bone 41, 794–803.
Marston, H.R., 1970. The requirement of sheep for cobalt or for vitamin B12. Br. J. Nutr. 24, 615–633. McDowell, L.R., 1985. Nutrition of Grazing Ruminants in Warm Climates. Academic Press, New York. McDowell, L.R., 1996. Feeding minerals to cattle on pasture. Anim. Feed Sci. Technol. 60, 247–271. McDowell, L.R., 2003. Minerals in Animal and Human Nutrition, 2nd ed. Elsevier, Amsterdam. McNaughton, S.J., 1988. Mineral nutrition and spatial concentrations of African ungulates. Nature 334, 343–345. McNaughton, S.J., 1990. Mineral nutrition and seasonal movements of African migratory ungulates. Nature 345, 613–615. Muir, P.D., Sykes, A.R., Barrell, G.K., 1987. Calcium metabolism in red deer (Cervus elaphus) offered herbages during antlerogenesis: kinetic and stable balance studies. J. Agric. Sci. 109, 357–364. Muller, L.D., Schaffer, L.V., Ham, L.C., Owens, M.J., 1977. Cafeteria-style freechoice mineral feeder for lactating dairy cows. J. Dairy Sci. 60, 1574–1582. Richter, C.P., 1943. Total self-regulatory functions in animals and human beings. Harvey Lect. 38, 63–103. Schwarz, F.J., Kirchgessner, M., Stangl, G.I., 2000. Cobalt requirement of beef cattle-feed intake and growth at different levels of cobalt supply. J. Anim. Physiol. Anim. Nutr. 83, 121–131. Seagle, S.W., McNaughton, S.J., 1992. Spatial variation in forage nutrient concentrations and the distribution of Serengeti grazing ungulates. Landsc. Ecol. 70, 229–241. Smith, R.M., Marston, H.R., 1970. Some metabolic aspects of vitamin B12 deficiency in sheep. Br. J. Nutr. 24, 879–891. Stangl, G.I., Schwarz, F.J., Kirchgessner, M., 1999. Moderate longterm cobaltdeficiency affects liver, brain and erythrocyte lipids and lipoproteins of cattle. Nutr. Res. 19, 415–427. Stephens, D.W., Krebs, J.R., 1986. Foraging Theory. Princeton University Press, Princeton. Tracy, B.F., McNaughton, S.J., 1995. Elemental analysis of mineral lick soils from the Serengeti National Park, the Konza Prairie and Yellowstone National Park. Ecography 18, 91–94. Villalba, J.J., Provenza, F.D., Hall, J.O., 2008. Learned appetites for calcium, phosphorus, and sodium in sheep. J. Anim. Sci. 86, 738–747. Ward, M., Lardy, G., 2005. Beef Cattle Mineral Nutrition. AS-1287. NDSU Extension Service. North Dakota State University. Weeks, J.S., Kirkpatrick, C.M., 1976. Adaptations of white-tailed deer to naturally occurring sodium deficiencies. J. Wildl. Manage. 40, 610–625.
