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Energy supplementation as strategy of pasture management

ABSTRACT.

This study evaluated the effect of increased energy via supplementation on the performance, ingestive behavior, nutrient digestibility, and nitrogen metabolism of grazing heifers fed tropical forage in the rainy-dry transition season. Treatments consisted of mineral supplementation ad libitum (control) and multiple supplements formulated to provide different energy levels and the same amount of protein (300 g CP animal d-1) and were denominated as low (LE; 340 g TDN animal d-1), medium (ME; 780 g TDN animal d-1) and high (HE; 1220 g TDN animal d-1) energy. Animals supplemented with ME, and HE had a greater average daily gain in relation to the control treatment, with an increase of 41 and 46%, respectively. Greater values for total apparent digestibility of neutral detergent fiber were observed for the treatment HE. Lesser values of urinary urea N were observed for the control and HE treatments. Our results define the use of energy levels in the supplement as a tool for pasture management. If the purpose of the production system is to enhance forage intake, the option is to supply supplements with less energy levels. In contrast, if the purpose is to increase the stocking rate, supplements with greater energy levels should be used.

Keywords:
beef cattle; multiple supplementation; nutritional parameters; tropical pastures

Introduction

Tropical pastures are the basis of Brazilian beef cattle feed. But even pastures fertilized during a period of high rainfall does not constitute a complete food for the herd, and therefore, its association with correct mineral supplementation is the premise for adequate animal development. In these production systems, the insertion of protein and energy via supplement is performed to overcome the nutritional deficits that depend on the forage characteristics, animal category, and desired productive performance.

The use of supplementation for grazing animals also constitutes a pasture management strategy, with changes in the stocking rate (Barbero et al., 2015Barbero, R. P., Malheiros, E. B., Araújo, T. L. R., Nave, R. L. G., Mulliniks, J. T., Berchielli, T. T., ... Reis, R. A. (2015). Combining Marandu grass grazing height and supplementation level to optimize growth and productivity of yearling bulls. Animal Feed Science and Technology, 209(1), 110-118. DOI: https://doi.org/10.1016/j.anifeedsci.2015.09.010
https://doi.org/https://doi.org/10.1016/...
; Fajardo et al., 2015Fajardo, N. M., Poli, C. H. E. C., Bremm, C., Tontini, J. F., Castilhos, Z. M. S., McManus, C. M., ... Monteiro, A. L. G. (2015). Effect of concentrate supplementation on performance and ingestive behaviour of lambs grazing tropical Aruana grass (Panicum maximum). Animal Production Science, 56(10), 1693-1699. DOI: https://doi.org/10.1071/AN14698
https://doi.org/https://doi.org/10.1071/...
), due to the associative effect of the supplement with the forage that modifies the ruminal metabolic condition (Dixon & Stockdale, 1999Dixon, R. M., & Stockdale, C. R. (1999). Associative effects between forages and grains: Consequences for feed utilisation. Australian Journal of Agricultural Research, 50(5), 757-773. ). Additionally, as there is a reduction in the amount of rainfall, a period recognized as a rainy-dry transition, there is a reduction in the quantity and quality of the forage produced, with emphasis on the beginning of the decrease in the nitrogen concentration.

In this situation, there is disagreement among some authors on the protein and energy levels that should be provided by the supplement to optimize the pasture use, since several works (Cabral, Paulino, Paula, Valadares, & Araújo, 2012Cabral, C. H. A., Paulino, M. F., Paula, N. F., Valadares, R. F. D., & Araújo, F. L. (2012). Levels of supplementation for grazing pregnant beef cows during the dry season. Revista Brasileira de Zootecnia, 41(12), 2441-2449.; Cabral et al., 2014aCabral, C. H. A., Paulino, M. F., Detmann, E., Valente, É. E. L., Barros, L. V., & Cabral, C. E. A. (2014a). Levels of supplementation for grazing pregnant beef heifers. Bioscience Journal, 30(1), 226-234. ; Cabral et al., 2014bCabral, C. H. A., Paulino, M. F., Detmann, E., Valadares Filho, S. C., Barros, L. V., Valente, É. E. L., ... Cabral, C. E. A. (2014b). Levels of supplementation for grazing beef heifers. Asian-Australasian Journal of Animal Sciences, 27(6), 806-817. DOI: https://doi.org/10.5713/ajas.2013.13542
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) reported that increasing supplement supply without adequate protein and energy levels promotes an imbalance of nutrients and impairs animal performance.

Some authors claim that due to the still expressive protein concentration in this period, supplementation with a greater energy level should be promoted (Sales et al., 2008Sales, M. F. L., Paulino, M. F., Porto, M. O., Valadares Filho, S. C., Acedo, T. S., & Couto, V. R. M. (2008). Energy levels in multiple supplements for finishing beef cattle grazing palisade grass pasture during the rainy to dry transition season. Revista Brasileira de Zootecnia, 37(9), 1704-1712. DOI: https://doi.org/10.1590/S1516-35982008000900025
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). In opposition, Porto et al. (2011Porto, M. O., Paulino, M. F., Detmann, E., Valadares Filho, S. C., Sales, M. F. L., Cavali, J., ... Acedo, T. S. (2011). Offers of multiple supplements to crossbreds Nellore young bulls in the growing stage on pasture, during the dry season: productive performance and nutritional characteristics. Revista Brasileira de Zootecnia, 40(11), 2548-2557. DOI: https://doi.org/10.1590/s1516-35982011001100037
https://doi.org/https://doi.org/10.1590/...
) and Santos et al. (2019Santos, A. R. M., Cabral, C. H. A., Cabral, C. E. A., Barros, L. V, Barros, J. M., Cabral, W. B., & Dias, M. R. (2019). Energy to protein ratios in supplements for grazing heifers in the rainy season. Tropical Animal Health and Production, 51(8), 2395-2403. DOI: https://doi.org/10.1007/s11250-019-01953-8
https://doi.org/https://doi.org/10.1007/...
) did not observe an increase in weight gain with the increase of energy insertion in the supplement.

Complementarily, Detmann, Paulino, Valadares Filho, and Huhtanen (2014Detmann, E., Paulino, M. F., Valadares Filho, S. C., & Huhtanen, P. (2014). Nutritional aspects applied to grazing cattle in the tropics: A review based on Brazilian results. Semina: Ciências Agrárias, 35(4), 2829-2854. DOI: https://doi.org/10.5433/1679-0359.2014v35n4Suplp2829
https://doi.org/https://doi.org/10.5433/...
)analyzed forage harvested from tropical pastures under continuous management and observed that most of the samples presented an energy and protein ratio above those demanded by the animals. In this situation, animals seek to reduce discomfort due to excess energy in the diet, alter the use of fiber, and thus reduce consumption (Forbes, 2003Forbes, J. M. T. (2003). The multifactorial nature of food intake control. Journal of Animal Science, 81(14,suppl 2), 139-144. DOI: https://doi.org/10.2527/2003.8114_suppl_2E139x
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).

Based on this, we hypothesize that the greatest impact of supplementation with greater energy levels will be on weight gain per area and, therefore, the objective wasto evaluate the effect of increased energy via supplementation on the performance, ingestive behavior, nutrient digestibility, and nitrogen metabolism of grazing heifers fed tropical forage.

Material and methods

Experimental site, animals, and management

The experiment was carried out in Rondonópolis-MT, Brazil (16° 28'S, 54°31' O), during the rainy-dry transition season, from 11 March to 31 May 2015. Air temperature data (°C) during the experimental period was obtained from a recording station of the Federal University of Mato Grosso- Rondonópolis Campus and the monthly precipitation was measured with a rain gauge located in the experimental area (Figure 1).

Forty crossbred heifers (Nelore breed predominance), with average initial ages and weights of 17 months and 229 kg, respectively, were assigned to an experimental area of 7 ha, consisting of four 1.75-ha paddocks uniformly covered with Marandu palisadegrass [Urochloa brizantha (Hochst. ex A. Rich.) R. D. Webster]. The animals were managed and cared for according to the guidelines and recommendations of the International Guiding Principles for Biomedical Research Involving Animals (CIOMS) - Geneva, 1985. All procedures were approved by the Federal University of Mato Grosso (register no. 63/2014), Rondonópolis, Brazil.

Figure 1
Rainfall and temperature during the experimental period.

Based on soil analyses and as recommended by Martha Júnior et al. (2007Martha Júnior, G. B., Vilela, L., & Sousa, D. M. G. (2007). Cerrado: uso eficiente de corretivos e fertilizantes em pastagens. Planaltina, DF: Embrapa Cerrados.), fertilization with urea (100 kg ha-1) and potassium chloride (50 kg ha-1), was applied to all paddocks, not requiring liming and phosphate fertilization. Fertilizers were applied on 12 December 2014, 2 January 2015, and 20 March 2015.

Treatments (Table 1) consisted of mineral supplementation ad libitum (control) and multiple supplements formulated to provide different energy levels and the same amount of protein (300 g CP animal d-1) and were denominated as low (LE; 340 g TDN animal d-1), medium (ME; 780 g TDN animal d-1) and high (HE; 1220 g TDN animal d-1) energy supplements. Treatments LE, ME, and HE were provided in the amount of 0.5, 1.0, and 1.5 kg animal d-1, respectively, daily between 10h00 and 12h00, in covered troughs, allowing access by the animals on both sides. Animals had unrestricted access to water throughout the experiment.

Productive performance

Heifers were weighed at the beginning and end of the experiment, after a 14-h fasting period. Total weight gain (TWG) was determined as the difference between the final and initial fasting body weight, and the average daily gain (ADG) was quantified as the ratio between TWG and the number of experimental days (84 days).

Table 1
Proportion of ingredients (based on natural matter) and chemical composition (based on dry matter) of supplements and ingredients amount supply.

Ingestive behavior

Ingestive behavior was determined on days 31 and 52 of the experiment for 12 consecutive hours, from 06h00 to 18h00. Grazing time, rumination time, and idle time were evaluated every 15 minutes. Time spent in forage selection, apprehension, and semi-digested food manipulation, which included the short time intervals used for the forage selection, was denominated grazing time. For rumination time, we considered the time spent in regurgitation and semi-digested food remastication, and the time elapsed between swallowing and regurgitation. When the animals showed no locomotive activity and an absence of mandibular movements, this was considered idle time (Cabral, Bauer, Cabral, Souza, & Benez, 2011Cabral, C. H. A., Bauer, M. O., Cabral, C. E. A., Souza, A. L., & Benez, F. M. (2011). Diurnal ingestive behavior of steers supplemented at rainy season. Revista Caatinga, 24(4), 178-185.).

Feed efficiency for DMI (FEDM) and NDF intake (FENDF); and rumination efficiency for DMI (REDM) and NDF intake (RENDF) were obtained by the following equations: FEDM = DMI/GT; FENDF = NDFI/GT; REDM = DMI/RT; RENDF = NDFI/RT.

Where: DMI, dry matter intake; NDFI, neutral detergent fiber intake; GT, grazing time; RT, rumination time.

Experimental procedures and sampling

Forage samples were collected monthly at representative points of the mean canopy height with the use of a metal frame (0.25 m2). Average canopy height was obtained from 100 points per paddock. All forage samples were collected at ground level. After collection, a sub-sample was taken to the laboratory for manual separation of the morphological components: leaf blade, stem + sheath, and dead material. These samples were oven-dried at 55°C for 72 hours and subsequently weighed to obtain dry weights. Also, forage samples for chemical analysis were collected monthly using the hand-plucked technique (Sollenberger & Cherney, 1995Sollenberger, L. E., & Cherney, D. (1995). Evaluating forage production and quality. In R. F. Barnes, & D. Miller (Eds.), Forages: the science of grassland agriculture (p. 97-110). Ames, IA: Iowa State University Press.) and oven-dried at 55°C for 72 hours and ground to 1 and 2 mm in a knife mill.

To evaluate the intake and digestibility of the diet components, a digestibility trial was conducted over nine days, starting on the 35th and finishing on the 43rd day of the productive performance evaluation. The first six days were used for the adaptation of animals to the markers and the last three days were used for the collection of feces and urine.

Chromium oxide (Cr2O3) was used as an external maker to estimate fecal excretion and was supplied daily at 09h00 to each animal, at a dose of 15 g, introduced orally directly into the esophagus. Individual intake of supplement was estimated using titanium dioxide (TiO2), being 15 g of TiO2 per animal added to the supplement.

During the last three days of the trial, fecal samples were collected at different times for each collection day: 15h00, 10h00, and 7h00 hours, respectively. Approximately 200 g of feces were collected per animal immediately after defecation or directly from the rectum. The samples were packed in plastic bags, individually labelled, and oven-dried dried at 55ºC. After drying, the samples were ground in a knife mill (1-mm sieve); samples were consolidated per animal, referring to the 3 days of collection (Table 2).

Table 2
Morphological and chemical composition of Marandu palisadegrass.

Chemical analysis

Forage and supplement samples were analyzed for dry matter (DM), mineral matter (MM), crude protein (CP; Silva & Queiroz, 2002Silva, D. J., & Queiroz, A. C. (2002). Análise de alimentos: métodos químicos e biológicos. Viçosa, MG: UFV.) and neutral detergent fiber (NDF; Mertens, 2002Mertens, D. R. (2002). Gravimetric determination of amylase-treated neutral detergent fiber in feeds with refluxing in beakers or crucibles: collaborative study. Journal of AOAC International, 85(6), 1217-1240.), using thermostable alpha-amylase and omitting the use of sodium sulfite; neutral detergent insoluble fiber (NDFI; Valente et al., 2011Valente, T. N. P., Detmann, E., Valadares Filho, S. C., Cunha, M., Queiroz, A. C., & Sampaio, C. B. (2011). In situ estimation of indigestible compounds contents in cattle feed and feces using bags made from different textiles. Revista Brasileira de Zootecnia, 40(3), 666-675. DOI: https://doi.org/10.1016/j.nima.2015.01.075
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) were quantified by in situ incubation procedures with Ankon® bags (F57) for 288 hours in samples processed at 2-mm, and acid detergent insoluble protein (ADIP) was determined according to the methodology described by Licitra, Hernandez, and Van Soest (1996Licitra, G., Hernandez, T. M., & Van Soest, P. J. (1996). Standardization of procedures for nitrogen fractionation of ruminant feeds. Animal Feed Science Technology, 57(1), 347-358. DOI: https://doi.org/https://doi.org/10.1016/0377-8401(95)00837-3
https://doi.org/https://doi.org/https://...
).

Fecal samples were evaluated for TiO2 content, according to the colorimetric technique described by Titgemeyer, Armendariz, Bindel, Greenwood, and Löest (2001Titgemeyer, E. C., Armendariz, C. K., Bindel, D. J., Greenwood, R. H., & Löest, C. A. (2001). Evaluation of titanium dioxide as a digestibility marker for cattle. Journal of Animal Science, 79(4), 1059-1063. DOI: https://doi.org/10.2527/2001.7941059x
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). Chromium oxide content was evaluated in an atomic absorption spectrophotometer, according to Williams, David, and Iismaa (1962Williams, C. H., David, D. J., & Iismaa, O. (1962). The determination of chromic oxide in faeces samples by atomic absorption spectrophotometry. The Journal of Agricultural Science, 59(3), 381-385. DOI: https://doi.org/10.1017/S002185960001546X
https://doi.org/https://doi.org/10.1017/...
). Fecal excretion was estimated by an intermediate of the ratio between the provided dose of the indicator and the fecal concentration of chromium oxide (Smith & Reid, 1955Smith, A. M., & Reid, J. T. (1955). Use of chromic oxide as an indicator of fecal output for the purpose of determining the intake of pasture herbage by grazing cows. Journal of Dairy Science, 38(5), 515-524.)

To estimate voluntary forage intake, the internal iNDF indicator was used, according to Detmann et al. (2001Detmann, E., Paulino, M. F., Zervoudakis, J. T., Valadares Filho, S. C., Euclydes, R. F., Lana, R. P., & Queiroz, D. S. (2001). Chromium and internal markers in intake determination by crossbred steers, supplemented at pasture. Revista Brasileira de Zootecnia, 30(5), 1600-1609. DOI: https://doi.org/10.1590/S1516-35982001000600030
https://doi.org/https://doi.org/10.1590/...
). An estimate of the individual supplement intake was obtained according to Santos et al. (2019Santos, A. R. M., Cabral, C. H. A., Cabral, C. E. A., Barros, L. V, Barros, J. M., Cabral, W. B., & Dias, M. R. (2019). Energy to protein ratios in supplements for grazing heifers in the rainy season. Tropical Animal Health and Production, 51(8), 2395-2403. DOI: https://doi.org/10.1007/s11250-019-01953-8
https://doi.org/https://doi.org/10.1007/...
). Total dry matter intake (kg d-1) was estimated as the sum of individual forage dry matter intake and individual supplement intake.

Nitrogen metabolism

Spot urine samples from spontaneous urination were collected from all animals (Valadares et al., 1999Valadares, R. F. D., Broderick, G. A., Valadares Filho, S. C., & Clayton, M. K. (1999). Effect of replacing alfalfa silage with high moisture corn on ruminal protein synthesis estimated from excretion of total purine derivatives. Journal of Dairy Science, 82(12), 2686-2696. DOI: https://doi.org/10.3168/jds.S0022-0302(99)75525-6
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) on the 9th day of the digestibility trial, 4h after feeding. One sample of 10 mL was diluted in 40 mL of H2SO4 (0.036 N) to reduce the pH to values below 3.0, avoiding nitrogen loss. The concentrations of creatinine, urea, and purine derivates were determined in the diluted sample.

The creatinine content was determined according to the modified Jaffé method, while uric acid was measured using the enzymatic-colorimetric method with a clear lipid factor. Allantoin concentrations were determined according to the colorimetric method described by Chen and Gomes (1992Chen, X. B., & Gomes, M. J. (1992). Estimation of microbial protein supply to sheep and cattle based on urinary excretion of purine derivatives - an overview of the technical details. Aberdeen, UK: Rowett Research Institute. DOI: https://doi.org/10.1016/j.cvfa.2007.03.003
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), and urea was measured via the urease method/glutamate dehydrogenase (GLDH). Total urinary volume was estimated based on the relationship between daily creatinine excretion as a function of body weight and the creatinine concentration in urine. Creatinine excretion per unit of body weight was obtained via the equation in Costa e Silva et al. (2012Costa e Silva, L. F., Valadares Filho, S. C., Chizzotti, M. L., Rotta, P. P., Prados, L. F., Valadares, R. F. D., ... Braga, J. M. S. (2012). Creatinine excretion and relationship with body weight of Nellore cattle. Revista Brasileira de Zootecnia, 41(3), 807-810. DOI: https://doi.org/10.1590/S1516-35982012000300046
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).

Urinary urea N (UreaN) was estimated as the product between the concentration in the urine spot samples and the estimated urinary volume. The excretion of total purine derivatives (mmol d-1) was calculated as the sum of the amounts of allantoin, and uric acid excreted via the urine. The absorbed purines were calculated from the excretion of total purine derivatives, using the equation in Barbosa et al. (2011Barbosa, A. M., Valadares, R. F. D., Valadares Filho, S. C., Pina, D. S., Detmann, E., & Leão, M. I. (2011). Endogenous fraction and urinary recovery of purine derivatives obtained by different methods in Nellore cattle. Journal of Animal Science, 89(2), 510-519. DOI: https://doi.org/10.2527/jas.2009-2366
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). Ruminal synthesis of microbial nitrogen (NMIC) was estimated based on the absorbed purines, using the equation proposed by Chen and Gomes (1992Chen, X. B., & Gomes, M. J. (1992). Estimation of microbial protein supply to sheep and cattle based on urinary excretion of purine derivatives - an overview of the technical details. Aberdeen, UK: Rowett Research Institute. DOI: https://doi.org/10.1016/j.cvfa.2007.03.003
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) and NTOTAL in bacteria, which is 0.134 according to Valadares et al. (1999Valadares, R. F. D., Broderick, G. A., Valadares Filho, S. C., & Clayton, M. K. (1999). Effect of replacing alfalfa silage with high moisture corn on ruminal protein synthesis estimated from excretion of total purine derivatives. Journal of Dairy Science, 82(12), 2686-2696. DOI: https://doi.org/10.3168/jds.S0022-0302(99)75525-6
https://doi.org/https://doi.org/10.3168/...
).

Blood samples were collected via jugular venipuncture, using commercial vacuum kits and coagulation accelerator gel, and were immediately centrifuged at 400 rpm for 15 minutes. The obtained serum was frozen at -20°C for the quantification of serum urea nitrogen (SUN). Blood serum samples were analyzed for urea levels via the urease/GLDH method.

Experimental design and statistical analysis

The experimental design was a completely randomized design, with four treatments. Data were analyzed using the fixed-model method, with a parametric structure, using the SAS® statistical software. A multiple comparison of the means, associated with the fixed effect of supplementation, was performed with a Tukey test (p = 0.05). The mathematical model was as follows: Yij = μ + Di(j) + eij, where Yij was the dependent variable of the supplementation i, measured in animal j; μ was the overall mean; Di was the fixed effect of supplementation i, measured in animal j; and eij was the random error of supplementation i, measured in animal j.

Results

There was an effect of supplementation on the animals' performance (p < 0.05; Table 3). Animals supplemented with ME, and HE had a greater ADG in relation to the control treatment, with an increase of 41 and 46%, respectively. However, there was no difference between supplemented animals.

Supplementation affects the ingestive behavior (p < 0.05; Table 3). Lesser grazing times were observed for animals supplemented with HE. Animals of LE treatments had lesser rumination times, however, animals supplemented with HE had longer idle time. Lesser REDM was observed for animals supplemented with LE, and greater FENDF was observed for animals supplemented with HE.

Table 3
Effect of energy levels in supplements on performance and ingestive behavior of grazing heifers.

There was no supplementation effect on intakes of total DM, CP, NDF, and digested NDF (p > 0.05; Table 4). However, there was a reduction of 41% on forage DM intake of animals supplemented with HE in relation to the control (5.33 kg d-1). In terms of nutrient digestibility, supplementation affects the total apparent digestibility of NDF, with greater values for animals supplemented with HE (Table 4).

Table 4
Effect of energy levels in supplements on intake and total apparent digestibility of nutrients of grazing heifers.

Regarding the nitrogen metabolism, a supplementation effect was observed for UreaN and SUN (p < 0.05; Table 5). Lesser values of UreaN were observed for the control and HE treatments. Supplemented animals presented greater SUN values.

Table 5
Effect of energy levels in supplements on nitrogen metabolism of grazing heifers

Discussion

For better supplement use efficiency, it is necessary to know the plant and animal interface, which involves studies of how grazing conditions interfere with the ingestive behavior, intake, and performance, to identify the appropriate management conditions for the animal production system. According to previous studies, the responses related to animals raised in pastures are directly related to the nutritional levels of the diet (Mendes et al., 2015Mendes, F. B. L., Silva, R. R., Carvalho, G. G. P., Silva, F. F., Lins, T. O. J. D’A, Silva, A. L. N., ... Guimarães, J. O. (2015). Ingestive behavior of grazing steers fed increasing levels of concentrate supplementation with different crude protein contents. Tropical Animal Health and Production, 47(2), 423-428. DOI: https://doi.org/10.1007/s11250-014-0741-z
https://doi.org/https://doi.org/10.1007/...
; Brandão et al., 2017Brandão, R. K. C., Carvalho, G. G. P., Silva, R. R., Dias, D. L. S., Mendes, F. B. L., Lins, T. O. J. D’A., ... Araujo, M. L. G. M. L. (2017). Correlation between production performance and feeding behavior of steers on pasture during the rainy-dry transition period. Tropical Animal Health and Production, 50(1), 105-111. DOI: https://doi.org/10.1007/s11250-017-1408-3
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).

With the additional supply of nutrients via supplementation, the daily metabolic requirements are met more quickly (Brandão et al., 2016Brandão, R. K. C., Carvalho, G. G. P., Silva, R. R., Dias, D. L. S., Mendes, F. B. L., Lins, T. O. J. D’A., ... Rufino, L. M. A., & Tosto, M. S. L. (2016b). Comparison of protein and energy supplementation to mineral supplementation on feeding behavior of grazing cattle during the rainy to the dry season transition. Springer Plus, 5(1), 933-939. DOI: https://doi.org/10.1186/s40064-016-2603-9
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b). Despite the difference in control treatment performance for the supplemented animals (Table 3), ADG of control treatment was satisfactory, with greater values than those previously found in similar studies (Brandão et al., 2016aBrandão, R. K. C., Carvalho, G. G. P., Silva, R. R., Dias, D. L. S., Guimaraes, J. O., Lins, T. O. J. D’A., ... Rufino, L. M. A. (2016a). Performance of dairy steers supplemented pasture during the transition period water-dry. Journal of Animal and Plant Sciences, 26(6), 1582-1588. ; Brandão et al., 2017Brandão, R. K. C., Carvalho, G. G. P., Silva, R. R., Dias, D. L. S., Mendes, F. B. L., Lins, T. O. J. D’A., ... Araujo, M. L. G. M. L. (2017). Correlation between production performance and feeding behavior of steers on pasture during the rainy-dry transition period. Tropical Animal Health and Production, 50(1), 105-111. DOI: https://doi.org/10.1007/s11250-017-1408-3
https://doi.org/https://doi.org/10.1007/...
; Almeida et al., 2018Almeida, D. M., Marcondes, M. I., Rennó, L. N., Martins, L. S., Alberto, F., Villadiego, C., & Paulino, M. F. (2018). Soybean grain is a suitable replacement with soybean meal in multiple supplements for Nellore heifers grazing tropical pastures. Tropical Animal Health and Production, 50(8), 1843-1849. DOI: https://doi.org/https://10.1007/s11250-018-1630-7
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). This occurred because the forage consumed had a relatively good nutritive value, with high CP concentration (> 120 g kg-1; Table 2), above that recommended for ruminal microorganism maintenance (70 g kg-1; Sampaio et al., 2010Sampaio, C. B., Detmann, E., Paulino, M. F., Filho, S. C. V., Souza, M. A., Lazzarini, I., ... Queiroz, A. C. (2010). Intake and digestibility in cattle fed low-quality tropical forage and supplemented with nitrogenous compounds. Tropical Animal Health and Production, 42(1), 1471-1479. DOI: https://doi.org/10.1007/s11250-010-9581-7
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). Besides that, there was an increase in forage mass in April and May as a result of the fertilization carried out in March and the precipitation in the experimental period (Table 2; Figure 1). This increase in the forage mass provides two management strategies, which are the increase in the stocking rate during the period of greater precipitaion, or the possibility of starting the dry season with a greater forage mass since this is a bottleneck in forage-based systems.

Treatments ME and HE allowed greater ADG, however, there is a reduction in forage DM intake compared to control, that is, there was a substitution effect of pasture by supplement, having seen the equal DM intake among all treatments. This effect is characterized by the reduction in forage DM intake with an increase in supplement intake while maintaining the total DM intake (Minson, 1990Minson, D. J. (1990). Forage in ruminant nutrition. San Diego, CA: Academic Press.). As forage is the most economical source of nutrient supply to animals, the adoption of supplements should optimize the use of fodder resources, promoting minimal substitution. However, ME and HE treatments become a viable alternative in systems where the objective is to increase the stocking rate. This is the reality in properties where the high value of the land requires the intensification of cattle raising, with an increase in the offtake rate and working capital invested.

Thus, while fertilization provides an increase in the stocking rate without changing the individual performance of the animals (Delevatti et al., 2019Delevatti, L. M., Cardoso, A. S., Barbero, R. P., Leite, R. G., Romanzini, E. P., Ruggieri, A. C., & Reis, R. A. (2019). Effect of nitrogen application rate on yield, forage quality, and animal performance in a tropical pasture. Scientific Reports, 9(1), 7596. DOI: https://doi.org/10.1038/s41598-019-44138-x
https://doi.org/https://doi.org/10.1038/...
), energy supplementation increases the stocking rate and the average daily gain of the animals, boosting meat production per area.

The ingestion of forage and supplements causes modifications in the energy: protein ratio of the diet, one of the determinants of consumption (Illius & Jessop, 1996Illius, A. W., & Jessop, N. S. (1996). Metabolic constraints on voluntary intake in ruminants. Journal of Animal Science, 74(12), 3052-3062. DOI: https://doi.org/10.2527/1996.74123052x
https://doi.org/https://doi.org/10.2527/...
), and the animal can make adjustments in this ratio by increasing or reducing the use of forage. Excess energy in the diet causes a decrease in consumption (Forbes, 2003Forbes, J. M. T. (2003). The multifactorial nature of food intake control. Journal of Animal Science, 81(14,suppl 2), 139-144. DOI: https://doi.org/10.2527/2003.8114_suppl_2E139x
https://doi.org/https://doi.org/10.2527/...
), while protein excess causes an increase in hepatic energy expenditure and, consequently, a reduction in productive performance (Detmann, Paulino, Valadares Filho, & Huhtanen, 2014Detmann, E., Paulino, M. F., Valadares Filho, S. C., & Huhtanen, P. (2014). Nutritional aspects applied to grazing cattle in the tropics: A review based on Brazilian results. Semina: Ciências Agrárias, 35(4), 2829-2854. DOI: https://doi.org/10.5433/1679-0359.2014v35n4Suplp2829
https://doi.org/https://doi.org/10.5433/...
).

Ingestive behavior variables confirm the explanations described above. The animals of LE treatment had the longest grazing times (Table 3) since depending on the amount of supplement included in the diet, the increase in supplementation levels promotes a reduction in grazing time because of the substitutive effect of pasture intake per concentrate (Minson, 1990Minson, D. J. (1990). Forage in ruminant nutrition. San Diego, CA: Academic Press.), which was observed in the present study.

Our rumination efficiency (RE) values are above what is in the literature because in this work the behavior was evaluated for 12 hours, while the works that evaluated these efficiencies did the behavior for 24 hours (Araújo et al., 2021Araújo, F. L., Souza, K. A., Moura Santana, N., Santana, L. R. C., Silva, C. S., Oliveira, K. N., ... Bagaldo, A. R. (2021). Animal performance, ingestive behavior, and carcass characteristics of grazing-finished steers supplemented with castor bean (Ricinus communis L.) meal protein. Tropical Animal Health and Production, 53(2), 240. DOI: https://doi.org/10.1007/s11250-021-02673-8
https://doi.org/https://doi.org/10.1007/...
; Silva et al., 2021Silva, T. S., Araujo, G. G. L., Santos, E. M., Oliveira, J. S., Campos, F. S., Godoi, P. F. A., ... Turco, S. H. N. (2021). Water intake and ingestive behavior of sheep fed diets based on silages of cactus pear and tropical forages. Tropical Animal Health and Production, 53(2), 244. DOI: https://doi.org/10.1007/s11250-021-02686-3
https://doi.org/https://doi.org/10.1007/...
). Thus, there is an increase in the RE values, due to the shorter times of rumination with emphasis on the fact that the animals ruminate with greater intensity at night (Kilgour, 2012Kilgour, R. J. (2012). In pursuit of 'normal': a review of the behaviour of cattle at pasture. Applied Animal Behaviour Science, 138(1-2), 1-11. DOI: https://doi.org/10.1016/j.applanim.2011.12.002
https://doi.org/https://doi.org/10.1016/...
).

Microbial growth is potentiated by the combination of the availability of fermentable energy and degradable nitrogen in the rumen (Russel, O’Connor, Fox, Van Soest, & Sniffen, 1992Russel, J. B., O’Connor, J. D., Fox, D. G., Van Soest, P. J., & Sniffen, C. J. (1992). A net carbohydrate and protein system for evaluating cattle diets: I. Ruminal fermentation. Journal of Animal Science, 70(1), 3551-3561. DOI: https://doi.org//1992.70113551x
https://doi.org/https://doi.org//1992.70...
). With the increase in nitrogen and energy supply due to the multiple supplementation, ruminal cellulolytic bacterial activity also increases (Silva-Marques et al., 2015Silva-Marques, R. P., Zervoudakis, J. T., Hatamoto-Zervoudakis, L. K., Cabral, L. D. S., Alexandrino, E., Neto, A. J., ... Melo, A. C. B. (2015). Multiple supplements for beef heifers on pasture during the dry season: nutritional characteristics. Semina: Ciências Agrárias, 36(1), 509-524. DOI: https://doi.org/10.5433/1679-0359.2015v36n1p509
https://doi.org/https://doi.org/10.5433/...
), which explains the greater NDF digestibility in the HE treatment. Besides that, the inclusion of supplement and the reduction in forage DM intake also contributes to the increase of the NDF digestibility, because the NDF of supplement is more digestible than the NDF of forage.

Supplemented animals received the same amount of protein via supplement, explaining the similar values for NI, UN, and FeN. However, a lesser NI was expected for the control treatment, which was not observed due to the pasture CP concentration (Table 2) and the reduction in forage DM intake in the supplemented treatments.

The lesser energy level of the supplement (LE treatment) and the difference in the speed of use of the protein and energy fractions of the supplement and the pasture increased UreaN. This can be observed by the non-difference in the intake of digested NDF, which consequently increased UreaN (Table 5).

The metabolic responses to protein supplementation in ruminants can be determined via the SUN concentration. The high concentration of SUN is related to the inefficient use of dietary CP (Broderick & Clayton, 1997Broderick, G. A., & Clayton, M. K. (1997). A statistical evaluation of animal and nutritional factors influencing concentrations of milk urea nitrogen. Journal of Dairy Science, 80(11), 2964-2971. DOI: https://doi.org/10.3168/jds.s0022-0302(97)76262-3
https://doi.org/https://doi.org/10.3168/...
). Valadares et al. (1999Valadares, R. F. D., Broderick, G. A., Valadares Filho, S. C., & Clayton, M. K. (1999). Effect of replacing alfalfa silage with high moisture corn on ruminal protein synthesis estimated from excretion of total purine derivatives. Journal of Dairy Science, 82(12), 2686-2696. DOI: https://doi.org/10.3168/jds.S0022-0302(99)75525-6
https://doi.org/https://doi.org/10.3168/...
) have stated that SUN values of 14 to 16 mg dL-1 result in dietary protein loss. For all supplemented treatments, the SUN values were within this range, while for the control treatment, the value was below it.

Despite the high CP concentration, the forage presented a significant amount of ADIP (Table 2), compromising the nutritive value of the diet, as ADIP is the fraction of N which represents the unavailable protein contained in the cell walls, resistant to the action of ruminal microorganisms and non-digestible in the intestines of ruminants (Sniffen, O’Connor, Van Soest, Fox, & Russell, 1992Sniffen, C. J., O’Connor, J. D., Van Soest, P. J., Fox, D. G., & Russell, J. B. (1992). A net carbohydrate and protein system for evaluating cattle diets: III. Cattle requirements and diet adequacy. Journal of Animal Science, 70(11), 3578-3596. DOI: http://www.ncbi.nlm.nih.gov/pubmed/1334063
https://doi.org/http://www.ncbi.nlm.nih....
).

Conclusion

For production systems that wish to maximize the use of tropical forages, with a minimum of substitute effect from the concentrate, the energy protein supplement indicated, when forage has a crude protein concentration of up to ~120 g kg-1, is the low energy supplement (340 g TDN animal d-1), resulting in satisfactory weight gain and a high nitrogen use efficiency.

Our results define the use of energy levels in the supplement as a tool for pasture management. If the purpose of the production system is to stimulate the forage intake, the option is to supply supplements with less energy levels. In contrast, if the purpose is to increase the stocking rate, supplements with greater energy levels should be used.

Acknowledgements

This work was supported by the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) [Grant Award Number: 482892/2013-7]

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Publication Dates

  • Publication in this collection
    12 Aug 2022
  • Date of issue
    2022

History

  • Received
    12 Sept 2020
  • Accepted
    05 Oct 2021
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