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Downloaded from https://academic.oup.com/tas/advance-article-abstract/doi/10.1093/tas/txz004/5304791 by Ed 'DeepDyve' Gillespie user on 05 February 2019 Kimberly M. Whittet, Andrea K. Watson, Galen E. Erickson, and Terry J. Klopfenstein Department of Animal Science, University of Nebraska, Lincoln, NE 68583-0908 ABSTRACT: A series of total urine collections urinary metabolites. Gestation did not affect UC were conducted to evaluate the effects of age, diet, (P = 0.42) or PD:C (P = 0.30). To evaluate the gestation, and body condition score (BCS) on uri- relationship between 12th rib fat thickness and nary creatinine (UC) and purine derivative (PD) metabolite excretion, 40 heifers were fed a com- excretion in heifers and cows. For each collection, mon finishing diet. There was no relationship urine was collected over a 5-d period and compos- between UC (mg/kg BW; P = 0.28) or PD:UC ited by animal within day. Daily samples were ana- (P = 0.47) and 12th rib fat thickness. To evaluate lyzed for UC and PD concentration and averaged the relationship between BCS and UC, 11 cows over the 5-d period. All animals were fed in indi- were fed a forage diet supplemented with DDG. vidual stanchions at 2.0% of body weight (BW). There was no relationship between BCS and UC To evaluate the relationship between age and UC (mg/kg BW; P = 0.99) or PD:C (P = 0.84). To excretion, 21 animals ranging from 5 to 80 months evaluate daily and diurnal variation in UC, nine of age were fed a forage-based diet supplemented heifers were fed a forage diet supplemented with with dried distillers grains (DDG). Creatinine DDG. Seven of the heifers were fed a finishing excretion (mg/kg BW) was not correlated with diet (90% concentrate, 10% forage) in a second age (P = 0.37). To determine if diet alters UC, 11 period. Urine was collected every 2 h from 0600 heifers were sampled for two urine collection peri- to 1800 hours. When expressed as mg/kg BW, UC ods. In period 1, heifers were fed a forage-based excretion was not different across animals fed the diet supplemented with DDG. In period 2, heifers forage-based (P = 0.40) or concentrate-based diet were fed a finishing diet (90% concentrate, 10% (P = 0.18). Stepwise regression indicated that at forage). Creatinine excretion (mg/kg BW) and least 3 d of collection were required to estimate PD:creatinine (PD:C) was greater (P = 0.01) for UC. Time within day and day within period effects heifers when fed the forage-based diet than when were observed (P < 0.01) for UC from 2-h interval fed the concentrate-based diet. Eleven cows fed a samples. The UC varies with type of diet and diur- forage-based diet supplemented with DDG were nal variation is present. Variation among animals sampled to determine the effect of gestation on is relatively small. Key words: beef cattle, creatinine excretion, purine derivatives, urine metabolites, urine output XX XX © The Author(s) 2019. Published by Oxford University Press on behalf of the American Society of XXXX Animal Science. All rights reserved. For permissions, please e-mail: firstname.lastname@example.org. This is an Open Access article distributed under the terms of the Creative Commons Attribution Non- Commercial License (http://creativecommons.org/licenses/by-nc/4.0/), which permits non-commer- cial re-use, distribution, and reproduction in any medium, provided the original work is properly cited. For commercial re-use, please contact email@example.com Transl. Anim. Sci. 2019.XX:XX–XX doi: 10.1093/tas/txz004 This project is based on research that was partially supported by the Nebraska Agricultural Experiment Sta- tion with funding from the Hatch Act (Accession Number Corresponding author: firstname.lastname@example.org 1007896) through the USDA National Institute of Food and Received October 17, 2018. Agriculture. Accepted January 24, 2019. 1 Downloaded from https://academic.oup.com/tas/advance-article-abstract/doi/10.1093/tas/txz004/5304791 by Ed 'DeepDyve' Gillespie user on 05 February 2019 2 Whittet et al. INTRODUCTION Table 1. Composition of experimental diets Ingredient Diet, % dry matter Measuring urine output in cattle is important Forage diet to allow research on N balance and metabolites Grass hay 70.0 in the urine such as creatinine and purine deriva- Dried distillers grains 30.0 tives (PD). Creatinine, a urine metabolite, has been Finishing diet shown to be an effective marker for estimating urine Dry-rolled corn 57.8 excretion in beef cattle (Chen et al., 1992; Valadares Wet corn gluten feed 30.0 et al., 1999; McDonald, 2003; Jardstedt et al., 2017). Alfalfa hay 10.0 A product of muscle metabolism, urinary creati- Limestone 1.8 nine (UC) excretion is directly related to muscle Sodium Chloride 0.3 mass (Lofgreen and Garrett, 1954; McCarthy et al., Trace mineral premix 0.05 1983; Gopinath and Kitts, 1984; Hayden et al., Vitamin A-D-E premix 0.02 1992) and is excreted at a constant rate relative to Rumensin premix 0.02 Tylan premix 0.01 BW (Brody, 1945). Swartz et al. (2016) and Brunsvig et al. (2017) used UC output to estimate N excre- Premix contained 10% Mg, 6% Zn, 2.5% Mn, 0.5% Cu, 0.3% I, and tion in grazing heifers and cows. Other researchers 0.05% Co. have used PD:creatinine (PD:C) ratio to estimate Premix contained 1,500 IU of vitamin A, 3,000 IU of vitamin D, and 3.7 IU of vitamin E per g. microbial crude protein (MCP) yield over a graz- Finishing diet contained 354 mg/kg monensin and 11 mg/kg tylosin ing season in heifers (MacDonald et al., 2007) and (dry matter basis; Elanco Animal Health, Greenfield, IN). cows (Patterson et al., 2006). Dórea et al. (2017), demonstrated a strong correlation between PD:C and both dry matter intake (DMI; R = 0.84) and on a blend of dry rolled corn and wet corn gluten digestible DMI (R = 0.85). Galyean and Tedeschi feed. All animals were fed at 2.0% of body weight (2014) further showed that MCP yield was highly (BW) once daily at 0800 hours and allowed ad libi- correlated to DMI (r = 0.88) and total digestible tum access to water. All animals were fed using indi- nutrients (TDN) intake (r = 0.89), which suggests vidual feed bunks within metabolism stanchions. a strong relationship between PD and MCP yield. Heifers and cows were housed in 1.3 × 2.5 m indi- It is important to understand the factors that vidual metabolism stanchions in a temperature-con- influence variability in UC excretion in beef cattle trolled room (25°C) and tethered for the 5 d collection to determine if it could be used as a reliable compo- period. Collection periods consisted of a minimum of nent in a system to predict urine output. The objec- 5 d for diet adaptation at ad libitum intake followed tives of these trials were to determine the effects of by 3 d of adaptation to restricted intake and then age, diet, gestation, body condition score (BCS), 5 d for sample collection at restricted intake (2.0% and 12th rib fat thickness on UC excretion and of BW). Animals were fitted with indwelling Foley PD:C concentration and total daily excretion. urethral catheters (C.R. Bard, Inc., Covington, GA; size 12 to 24) approximately 6 h prior to collection. Urethral catheters were connected to tubing at each MATERIALS AND METHODS animal’s hip with Velcro tape. Tubing was suspended All cattle were managed in accordance with by a cable and pulley and led to a 2-liter overnight protocols approved by the University of Nebraska urine collection bag (C.R. Bard, Inc., Covington, Institutional Animal Care and Use Committee. GA) in a cooled thermally-insulated container. Animals and Diets Trial 1. Twenty-one heifers and cows (BW range = 98 to 582 kg) were sampled to test the rela- Crossbred heifers and cows, (MARC III compos- tionship between age and UC and PD:C. Animals ite breed [¼ Angus, ¼ Hereford, ¼ Pinzgauer, and ¼ ranged from 5 to 80 mo of age and were fed the grass Red Poll]) were used in a series of total urine collec- hay diet supplemented with DDG. Age was approxi- tions to determine the effects of age, diet, gestation, mated to the nearest month at the time of collection. BCS, and 12th rib fat thickness on UC and PD:C. Animals were fed a diet (Table 1) of either grass hay Trial 2. To determine if diet alters UC and PD:C, (13% crude protein [CP]) supplemented with 30% heifers were sampled for two urine collection peri- dried distillers grains (DDG; 30% CP) or a common ods. In period 1, 10 heifers (BW = 404 kg, SD = 25) finishing diet ([16% CP; 77% dry matter [DM]) based were fed the grass hay diet supplemented with DDG. Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/advance-article-abstract/doi/10.1093/tas/txz004/5304791 by Ed 'DeepDyve' Gillespie user on 05 February 2019 Urinary creatinine in heifers and cows 3 In period 2, eight of the heifers (BW = 489 kg, Urine was collected continuously for 5 consec- SD = 25) were then fed the common finishing diet utive days. Urine was drained from each animal’s containing 87.8% concentrate (dry rolled corn and bag at 2-h intervals from 0600 to 1800 hours. Urine wet corn gluten feed), 10% forage (alfalfa hay), and was allowed to accumulate from 1800 to 0600 hours 2.2% supplement (fine ground corn carrier with when it was then drained. Drainage was measured in limestone, trace minerals, and Vitamin ADE). a 2 liters graduated cylinder to the nearest 10 ml and recorded. Approximately 45 ml aliquots of urine Trial 3. Five gestating (BW = 611 kg, SD = 42) were collected and stored in 50-ml screw-cap vials. and six non-gestating (BW = 531 kg, SD = 40) cows Aliquots were then composited by animal within day fed the grass hay diet supplemented with DDG by combining 1.0% of each collection interval (over- were sampled to determine the effect of gestation night sample and 2-h interval samples). Composited on UC and PD:C. Gestation was determined by daily samples and 2-h interval samples were then rectal palpation at the time of collection. initially diluted with 9-part urine diluent (Shingfield and Offer, 1999b) and 1-part urine. All samples were Trial 4. To evaluate the relationship between stored at –20°C until subsequent analysis. 12th rib fat thickness and UC and PD:C, heifers (n = 40; BW = 525 kg, SD = 65) were sampled at Laboratory Analyses and Calculations two stages during the finishing period. Heifers were Laboratory DM of feed ingredients and feed sampled in blocks of 10 animals and sampling peri- refusals were determined in a 100°C oven for 24 h. ods were approximately 75 d apart. Heifers were Crude protein was analyzed by the combustion fed the finishing diet containing 87.8% concentrate method (FP-528; LECO Corporation, St. Joseph, and 10% forage for both periods. Ultrasound 12th MI). Diluted urine samples were additionally rib fat thickness measurements were made between diluted with 4-parts urine diluent and 1-part diluted the 12th and 13th rib (Aloka 500V; Corometrics urine for analysis. Creatinine and PD (allantoin Medical Systems, Wallingford, CT). Measured 12th plus uric acid) concentrations were determined by rib fat thickness ranged from 0.8 to 2.4 cm. high-performance liquid chromatography (Waters Eleven cows (BW = 567 kg, SD = 57) were sam- Corp., Milford, MA) according to the procedure of pled to evaluate the relationship between BCS and Shingfield and Offer (1999b). UC and PD:C. Cows were fed the grass hay diet Daily excretion of creatinine (g/d) and PDs supplemented with DDG. Cow BCS was deter- (mmol/d) were calculated for each animal by multi- mined at the time of collection. Measured BCS plying the concentration (mmol/liter) by measured ranged from 3.9 to 6.4. 24 h urine volume. Creatinine was also expressed as a coefficient of BW (mg/kg BW) by dividing the Trial 5. Nine crossbred heifers (BW = 403 kg, daily creatinine excretion by each animal’s BW. SD = 27) were sampled for two total urine collec- tion periods to determine diurnal (patterns within a 24-h period), daily (comparison between consec- Statistical Analyses utive days), and animal variation in UC and PD:C in Regression analyses using the REG procedure growing and finishing heifers. In period 1, nine heif- of SAS (SAS Inst. Inc., Cary NC) were conducted ers were fed the forage-based diet supplemented with to evaluate the relationship between age, BCS, or DDG. In period 2, seven of the heifers (BW = 494 kg, 12th rib fat thickness and UC and PD:C (trials 1 SD = 22) were fed the common finishing diet con- and 4). The MIXED procedure of SAS was used taining 87.8% concentrate and 10% forage. to determine the effect of diet and gestation on UC and PD:C with animal included as a random effect Sampling (trials 2 and 3). For trial 5, data were analyzed using For all trials, experimental diets were sampled the MIXED procedure of SAS with day as a fixed once before each collection period and dried in a 60°C effect and animal as a random effect. Orthogonal forced-air drying oven for 48 h to determine DM linear, quadratic, and cubic effects within day content. Feed refusals were weighed, sampled, and were tested. Additionally, the STEPWISE and recorded daily before feeding and used to adjust DM RSQUARE selection methods of the REG pro- offered. Individual feed ingredients were sampled cedure of SAS were used to evaluate minimum before each collection period and dried at 60°C for number of days required for collection. Data were 48 h and then ground to pass through a 1-mm screen. compiled as periods consisting of day 1, 1 to 2, 1 to Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/advance-article-abstract/doi/10.1093/tas/txz004/5304791 by Ed 'DeepDyve' Gillespie user on 05 February 2019 4 Whittet et al. 0.75 3, and 1 to 4 and then regressed against the average metabolic BW (BW ). However, Shingfield and of days 1 through 5. In all trials, effects were con- Offer (1999a) suggest that tissue nucleic acid turno- sidered significant when P ≤ 0.05 and tendencies are ver is associated with an animal’s metabolic activity. discussed when P ≤ 0.10. If metabolic rate decreases as an animal ages, then the decrease observed could be accounted for by RESULTS AND DISCUSSION endogenous PD excretion. Trial 1 Trial 2 Regression analysis demonstrated that age was Excretion of UC (g/d) tended to be greater significantly correlated (r = 0.42) with UC excretion (P = 0.07) when heifers were fed the finishing diet (g/d). Because BW was correlated (r = 0.58) with compared to when they were fed the forage-based age, when UC excretion was expressed as a coef-fi diet (12.92 vs. 12.01 g/d; Table 2). However, when cient of BW (mg/kg BW), there was no correlation expressed as mg/kg BW, UC excretion was greater (r = 0.04; P = 0.37) between age and UC excretion (P = 0.01) for heifers fed the grass hay diet supple- (Figure 1). Range of UC excretion across all ani- mented with DDG than when fed the finishing diet. mals was 23.07 to 33.88 mg/kg BW with a mean When fed the forage-based diet, heifers excreted an of 28.64 mg/kg BW. It has been well documented average of 29.70 mg/kg BW compared to 26.50 mg/ that UC excretion is directly correlated with fat-free kg BW of UC when fed the finishing diet. The PD:C mass in ruminant animals (Lofgreen and Garrett, was greater (P < 0.01) for heifers fed the grass hay 1954; Van Niekerk et al., 1963; Hayden et al., 1992). diet supplemented with DDG than when fed the Additionally, Gopinath and Kitts (1984) showed an finishing diet. The PD:C ratio is potentially con- increase in UC excretion (g/d) in growing beef steers founded between age and diet. The range in ages with time indicating increased muscle mass with age. in Figure 1 is nearly 80 mo. The difference in age The ratio of molar concentrations of PD and of the heifers in this trial is less than 3 months so it UC decreased (r = 0.44) with age. The slope of would have very little effect on the values. −0.01 and intercept of 1.31 were both different Previous research indicates independence of (P < 0.01) from zero (Figure 1). PDs excreted in the UC excretion and level of dietary protein (Dinning urine are primarily exogenous in origin. However, et al., 1949; Butcher and Harris, 1957; Albin and due to tissue ATP and nucleic acid turnover, Clanton, 1966; Ørskov and MacLeod, 1982; endogenous PD excretion contributes to total PD Jardstedt et al., 2017) or energy intake (Van Niekerk excretion. Therefore, when calculating MCP, a cor- et al., 1963; Albin and Clanton, 1966). Diets uti- rection is made for contribution of endogenous PD lized in previous experiments were typically a base to the total supply of PD in the urine based upon diet with added levels of protein or energy intake. Conversely, the two diets fed in the current study 4.0 were formulated to be quite different. The grass hay 35 3.5 y = -0.03x + 29.30 diet supplemented with DDG would be expected to r = 0.04 3.0 give 0.4 kg/d of gain, while the finishing diet would 25 2.5 produce 1.5+ kg/d of gain. Furthermore, the for- age-based diet supplied fiber as the primary energy 2.0 15 1.5 y = -0.01x + 1.31 Table 2. Effect of diet (trial 2) and gestation (trial r = 0.44 1.0 3) on urinary creatinine and purine derivative excre- 5 0.5 tion in heifers (trial 2) and cows (trial 3) 0 0.0 0153045607590 Diet Forage Concentrate SEM P-value Age, months Creatinine, g/d 12.01 12.92 0.35 0.07 Creatinine, 29.70 26.50 0.86 0.01 Figure 1. Relationships between cattle age (months) and urinary mg/kg BW metabolites (n = 21; trial 1). Solid line and diamonds (left vertical PD:C molar ratio 1.27 0.95 0.03 <0.01 axis): relationship between age and urinary creatinine excretion (mg/ Gestation Gestating Non-gestating SEM P-value kg body weight). The slope of −0.03 (SE = 0.04) and intercept of 29.3 (SE = 1.16) were both different from zero (P < 0.01). Dashed line and Creatinine, g/d 17.67 14.59 0.76 0.01 squares (right vertical axis): relationship between age and the ratio of Creatinine, 29.20 27.67 1.35 0.42 molar concentrations of purine derivatives and creatinine. The slope mg/kg BW of −0.01 (SE = 0.002) and intercept of 1.31 (SE = 0.06) were both PD:C molar ratio 0.76 0.85 0.06 0.30 different from zero (P < 0.01). Translate basic science to industry innovation Creatinine, mg/kg BW Purine Derivative: Creatinine Downloaded from https://academic.oup.com/tas/advance-article-abstract/doi/10.1093/tas/txz004/5304791 by Ed 'DeepDyve' Gillespie user on 05 February 2019 Urinary creatinine in heifers and cows 5 source for the rumen microbes and the resulting approximately 246 to 277 d pregnant in the study end products of fermentation to the heifer. The fin- by Erb et al. (1977). Ørskov and MacLeod (1982) ishing diet was starch-based, producing a different reported no differences in UC excretion in relation microbial population and different end products. to BW in cows between 117 and 133 d pregnant and These two diets represent the extremes, yet the between 220 and 233 d pregnant. difference in UC excretion (mg/kg BW) was only There was no difference in PD:C (P = 0.30) 11.4%. Using the NRC (1996) model, the heifers between gestating and non-gestating cows (Table 2). were estimated to have 15% body fat when consum- The PD:C was numerically lower in gestating cows ing the forage-based diet. After the adaptation and compared to non-gestating cows (0.76 vs. 0.85) due treatment period, heifers were estimated to have to UC (g/d) excretion being numerically greater in 18% body fat. If 30% to 40% of the differences in gestating cows. UC excretion from different diets can be explained by greater percent body fat when the heifers were Trial 4 fed the finishing diet compared to the forage-based Ultrasound 12th rib fat thickness was poorly diet (NRC, 1996), then the impact of diet on UC correlated (r = 0.14) with UC excretion (g/d; excretion is small. The greater PD:C for heifers on data not shown). The slope of 1.76 and intercept the forage diet suggests greater microbial efficiency of 12.89 were both different from zero (P < 0.05). compared to the finishing diet. However, when expressed as a coefficient of BW (mg/kg BW; Figure 2), no relationship was found Trial 3 (r = 0.04; P = 0.28) between 12th rib fat thickness and creatinine output. The PD:C was not corre- The results of the effect of pregnancy on UC and lated (r = 0.02; P = 0.47) with 12th rib fat thick- PD:C in cows are shown in Table 2. Gestating cows ness. There was no relationship between BCS and had greater (P = 0.01) UC excretion (17.67 g/d) than UC excretion as mg/kg BW (r = 0.00; P = 0.99; cows that were not gestating (14.59 g/d). However, data not shown). Similarly, the PD:C was not cor- there was no difference (P = 0.42) in creatinine out- related with BCS (r = 0.005; P = 0.84; data not put when expressed as g/kg BW. Creatinine excre- shown). tion (mg/kg BW) for gestating and non-gestating UC excretion is highly correlated with empty cows was 29.20 and 27.67, respectively. Based on body protein and fat-free mass (Picón-Reátegui, Anthony et al. (1986), total conceptus weight would 1962; Van Niekerk et al., 1963). Therefore, it was range from 20 to 25 kg for the gestating cows. If we expected that ultrasound 12th rib fat thickness account for that weight (subtract it from cow body measurements and BCS could be used as predictors weight), it would increase the UC by 3% to 4%. However, it is not clear if the conceptus is a source 40 4.0 of creatinine. Because of these unknowns and the y = -1.27x + 30.52 35 3.5 small number of cows sampled, it might be inap- r = 0.04 propriate to conclude that gestating and non-ges- 30 3.0 tating cows produce similar amounts of UC. 25 2.5 Similar increases in UC excretion (g/d) were 20 2.0 observed by Erb et al. (1977) in dairy cows sampled y = 0.06x + 0.79 15 1.5 between 28 and 8 d prepartum and between 8 and r = 0.02 10 1.0 39 d postpartum. However, Erb et al. (1977) also reported differences in the creatinine coefficient 5 0.5 in cows sampled 28 and 8 d prepartum (24.96 mg/ 0 0.0 0.0 0.5 1.0 1.5 2.02.5 3.0 kg BW) and 8 and 39 d postpartum (21.12 mg/kg 12th rib backfat thickness, cm BW). Cows sampled in the current study were two groups of animals (gestating and non-gestating), Figure 2. Relationships between 12th rib fat thickness (cm) and uri- nary metabolites (n = 40; trial 4). Solid line and diamonds (left vertical not one group of animals sampled at two stages of axis): relationship between 12th rib fat thickness and urinary creatinine production, which would reduce animal variation. excretion (mg/kg body weight). The slope of −1.27 (SE = 1.15) was not Variation among individuals in the current study different from zero (P = 0.28) while the intercept of 30.52 (SE = 1.61) was different from zero (P < 0.01). Dashed line and squares (right ver- could account for the inability for a difference to be tical axis): relationship between 12th rib fat thickness and the ratio of detected between treatment groups. Additionally, molar concentrations of purine derivatives and creatinine. The slope cows sampled in this study ranged from approx- of 0.06 (SE = 0.08) was not different from zero (P = 0.47) while the imately 120 to 200 d pregnant compared to intercept of 0.79 (SE = 0.12) was different from zero (P < 0.01). Translate basic science to industry innovation Creatinine, mg/kg BW Purine Derivative: Creatinine Downloaded from https://academic.oup.com/tas/advance-article-abstract/doi/10.1093/tas/txz004/5304791 by Ed 'DeepDyve' Gillespie user on 05 February 2019 6 Whittet et al. of UC excretion or account for differences among forage diet (Table 4). Days 1 and 2 resulted in quad- animals due to fluctuations in body fat. However, no ratic (P ≤ 0.03) increases in PD:C over time, while relationships were found between ultrasound 12th day 4 responded cubically (P = 0.01). There was rib fat measurements or BCS and UC excretion. no effect (P ≥ 0.18) of time of collection on days 3 and 5 (P ≥ 0.18). There was not a significant day × time interaction (P = 0.35) for PD:C when heifers Trial 5 were fed the finishing diet (Table 4). Days 2 and 4 Excretion of UC expressed as mg/kg BW was resulted in linear (P = 0.001) increases in PD:C rel- constant across sampling days (P = 0.42), when ative to time of collection, while day 1 responded heifers were fed the forage diet. Range of UC excre- cubically (P = 0.02). There was no effect (P ≥ 0.12) tion was 28.11 to 30.28 mg/kg BW. The PD:C was of time of collection on days 3 and 5. not different by day (P = 0.17) across the 5 d period. Stepwise regression was used to evaluate the Excretion of UC expressed as mg/kg BW decreased minimum number of days required for an accurate linearly (P < 0.01) across experimental days when estimate of PD:C from average daily PD:C (Table heifers were fed the finishing diet. The PD:C was 4). The objective of stepwise regression was to eval- not different across collection days (P = 0.18) when uate a set of variables (days 1 through 5) that were heifers were fed the finishing diet. regressed against a dependent variable (average of days 1 through 5). Stepwise regression indicated Variation by day, time, and animal. Results for that 3 or 4 d of collection were required to accu- day and diurnal variation are represented in Tables rately estimate the PD:C when heifers were fed the 3 and 4 for UC excretion and PD:C, respectively. forage diet. Adjusted r values were 0.69 and 0.80 Diurnal variation was evaluated using 2-h inter- for 3 and 4 d, respectively. Collecting 4 d resulted in val samples from 0800 to 1800 hours for each ani- an 11 percentage unit increase in variance explained mal. There was a significant day × time interaction over 3 d of collection. However, 3 d resulted in a 23 (P < 0.01) for UC excretion (mg/kg BW) when heif- percentage unit increase in variance explained over ers were fed the forage diet (Table 3). Days 2 and 3 only 2 d of collection. Stepwise regression indicated had a linear increase in UC excretion (P ≤ 0.04) over that 2 or 3 d of collection were required when heif- time. On days 4 and 5, time of collection resulted ers were fed the finishing diet. Adjusted r values in a quadratic (P ≤ 0.01) increase in UC excretion. were 0.97 and 0.99 for 2 and 3 d, respectively. A significant (P < 0.01) day × time interaction was UC excretion was first noted for its constancy detected for UC excretion (mg/kg BW) when heifers when Folin began researching it in 1905. However, were fed the finishing diet (Table 3). Time of day since then, researchers have noted significant varia- resulted in a cubic response (P = 0.01) on days 1 and bility in creatinine output. Best et al. (1952) reported 2. Days 3, 4, and 5 resulted in quadratic (P ≤ 0.03) a significant diurnal cycle to creatinine excretion. increase in UC excretion over time of collection. However, Albin and Clanton (1966) found that N There was a significant day × time interaction was excreted in a similar diurnal cycle and when (P < 0.02) for the PD:C when heifers were fed the expressed as a ratio (N:creatinine), the variability Table 3. Effects of time within day on urinary creatinine excretion (mg/kg body weight) as predicted by 2-h interval samples (trial 5) Time of collection, hours Orthogonal contrasts Item 0800 1000 1200 1400 1600 1800 SEM Linear Quadratic Cubic Forage diet Day 1 33.92 25.32 34.31 22.26 35.75 29.43 2.32 0.87 0.17 0.14 Day 2 27.38 28.34 30.69 36.48 41.42 34.94 2.81 0.01 0.30 0.05 Day 3 21.02 30.27 34.65 23.28 39.11 31.75 3.87 0.04 0.30 0.48 Day 4 14.04 30.52 27.90 22.72 32.17 25.81 2.19 0.01 0.01 0.03 Day 5 19.03 28.41 30.21 28.40 28.29 24.18 1.95 0.16 0.01 0.20 Finishing diet Day 1 24.03 27.75 41.39 23.47 25.97 32.97 2.59 0.33 0.24 0.01 Day 2 23.43 26.15 42.22 24.00 27.49 33.53 2.44 0.08 0.14 0.01 Day 3 21.92 28.88 38.34 27.56 31.21 30.36 2.59 0.08 0.01 0.05 Day 4 25.22 21.50 39.13 16.22 25.87 18.96 2.49 0.04 0.03 0.35 Day 5 13.41 21.91 26.99 33.23 23.38 27.82 1.63 0.01 0.01 0.10 Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/advance-article-abstract/doi/10.1093/tas/txz004/5304791 by Ed 'DeepDyve' Gillespie user on 05 February 2019 Urinary creatinine in heifers and cows 7 Table 4. Effects of time within day on the purine derivative to creatinine molar ratio as predicted by 2-h interval samples (trial 5) Time of collection, hours Orthogonal contrasts Item 0800 1000 1200 1400 1600 1800 Mean SEM Linear Quadratic Cubic Forage diet Day 1 1.11 1.11 1.22 1.36 1.27 1.21 1.21 0.06 0.02 0.03 0.10 Day 2 1.10 0.99 1.20 1.10 0.91 0.83 1.02 0.06 0.01 0.01 0.70 Day 3 1.04 1.01 1.09 0.99 0.87 0.95 0.99 0.09 0.18 0.81 0.41 Day 4 1.07 1.01 1.21 1.35 1.38 1.17 1.20 0.08 0.01 0.03 0.01 Day 5 1.23 1.31 1.33 1.22 1.21 1.15 1.24 0.09 0.20 0.26 0.44 Finishing diet Day 1 1.20 1.17 1.20 1.29 1.35 1.36 1.26 0.05 0.01 0.14 0.02 Day 2 1.16 1.16 1.18 1.23 1.27 1.29 1.22 0.06 0.01 0.43 0.33 Day 3 1.10 1.06 1.04 1.04 1.09 1.11 1.07 0.06 0.67 0.12 0.75 Day 4 1.00 1.06 0.96 1.05 1.07 1.13 1.05 0.08 0.01 0.05 0.41 Day 5 1.13 1.11 1.11 1.11 1.11 1.12 1.12 0.07 0.91 0.41 0.80 40 2.5 was stabilized. The current data were analyzed to evaluate if there was a common trend in UC (mg/ 35 2.0 kg BW) excretion and PD:C across times of collec- tion within and across experimental days. If com- 1.5 mon trends were detected, then the variability in the excretion of each of these components could 1.0 be accounted for by collecting urine samples at spe- cific times of day. However, there were no common 0.5 trends in UC (mg/kg BW) excretion or PD:C. 0.0 The current results do demonstrate that there 12345678 9 was significant variation in UC excretion and PD:C Animal within and across days as predicted by 2-h inter- Figure 3. Animal variation in urinary creatinine excretion (mg/kg val samples. However, Butcher and Harris (1957) body weight) and the ratio of molar concentrations of PD:C while also found significant diurnal variation in creati- heifers were fed a hay diet supplemented with dried distillers grains (trial 5). Urinary creatinine excretion (mg/kg body weight) shown in nine and creatinine to N ratios, but when morning solid black bars following left vertical axis (SEM = 2.02; main effect and evening samples were composited, they had a of animal P = 0.40). Ratio of PD:C shown in gray bars following right similar ratio as total collections. Collecting multi- vertical axis (SEM = 0.08; main effect of animal P = 0.71). ple spot samples within a day may not be feasible in most production settings; however, collecting for multiple days may be practical. 40 2.5 When expressed as mg/kg BW, UC output was not different (P = 0.40; Figure 3) across experimen- 2.0 tal animals fed the forage diet. Mean UC excretion was 30.04 mg/kg BW. Additionally, the PD:C was 1.5 not different (P = 0.71) across experimental ani- 20 1.0 mals (Figure 3). When expressed as mg/kg BW, UC output was not different (P = 0.18) across animals 0.5 fed the finishing diet (Figure 4). Conversely, PD:C was different (P < 0.01) across animals fed the fin- 0.0 ishing diet (Figure 4). 1234567 Animal Research concerning the variation in creatinine excretion across animals is somewhat conflicting. Figure 4. Animal variation in urinary creatinine excretion (mg/kg When Folin (1905) first evaluated UC excretion, he body weight) and the ratio of molar concentrations of PD:C while heifers were fed a finishing diet (trial 5). Urinary creatinine excretion noted significant variation between individuals. More (mg/kg body weight) shown in solid black bars following left vertical recently, Jardstedt et al. (2017) reported no differ- axis (SEM = 1.97; main effect of animal P = 0.18). Ratio of PD:C ences in UC excretion due to diet (19.7 mg/kg BW), shown in gray bars following right vertical axis (SEM = 0.04; main effect of animal P = 0.002). but relatively high animal to animal variation (12.0 Translate basic science to industry innovation Creatinine, mg/kg BW Creatinine, mg/kg BW PD:C PD:C Downloaded from https://academic.oup.com/tas/advance-article-abstract/doi/10.1093/tas/txz004/5304791 by Ed 'DeepDyve' Gillespie user on 05 February 2019 8 Whittet et al. estrogen in cows prepartum and postpartum. J. Dairy Sci. to 32.5 mg/kg BW) using 48 h total urine collection. 60:1057–1063. doi:10.3168/jds.S0022-0302(77)83988-X Albin and Clanton (1966) reported significant animal Folin, O. 1905. Laws governing the chemical composition of variation in gestating cows but not in non-gestating urine. Am. J. Physiol. 13:66–115. cows. Excretion of UC (mg/kg BW) measured in this Galyean, M. L., and L. O. Tedeschi. 2014. Predicting microbial trial was relatively stable across experimental animals protein synthesis in beef cattle: relationship to intakes of total digestible nutrients and crude protein. J. Anim. Sci. with a mean excretion of 30.05 and 26.21 mg/kg BW 92:5099–5111. doi:10.2527/jas.2014-8098 for forage and finishing diets, respectively. Gopinath, R., and W. D. Kitts. 1984. Growth, N tau-methylhis- Influence of age, diet, and pregnancy on UC tidine excretion and muscle protein degradation in grow- excretion is minimal; therefore, creatinine has ing beef steers. J. Anim. Sci. 59:1262–1269. doi:10.2527/ promise as a marker of urine output and predic- jas1984.5951262x tion of PD excretion. Variation in excretion of both Hayden, J. M., W. G. Bergen, and R. A. Merkel. 1992. Skeletal muscle protein metabolism and serum growth hormone, UC and PD indicates that multiple days of collec- insulin, and cortisol concentrations in growing steers tion are required for an accurate estimate of PD:C. implanted with estradiol-17 beta, trenbolone acetate, or Further efforts in evaluating PD excretion should estradiol-17 beta plus trenbolone acetate. J. Anim. Sci. focus on relationships between spot urine samples 70:2109–2119. doi:10.2527/1992.7072109x and total collections and the ability of spot samples Jardstedt, M., A. Hessle, P. Nørgaard, W. Richardt, and E. Nadeau. 2017. Feed intake and urinary excretion collected on multiple days to estimate average PD of nitrogen and purine derivatives in pregnant suckler excretion. Variation in UC and PD excretion across cows fed alternative roughage-based diets. Livestock Sci. animals was minimal. 202:82–88. doi:10.1016/j.livsci.2017.05.026 Lofgreen, G. P., and W. N. Garrett. 1954. 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Translational Animal Science – Oxford University Press
Published: Jan 31, 2019
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