Effect of rate of body weight gain of steers during the stocker phase. II. Visceral organ mass and body composition of growing-finishing beef cattle

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1 Published January 28, 2015 Effect of rate of body weight gain of steers during the stocker phase. II. Visceral organ mass and body composition of growing-finishing beef cattle E. D. Sharman,* P. A. Lancaster,* C. P. McMurphy,* G. G. Mafi,* J. D. Starkey, C. R. Krehbiel,* and G. W. Horn* 1 *Oklahoma Agricultural Experiment Station, Stillwater 74078; and Texas Tech University, Lubbock ABSTRACT: Two experiments were conducted to examine the effect of rate of BW gain during the stocker phase on visceral organ mass and body composition of growing-finishing cattle that had grazed dormant native range (DNR) or winter wheat pasture (WP). In each experiment, fall-weaned steers were allotted randomly to 1 of these stocker production programs: 1) control, 1.02 kg steer 1 d 1 of a 40% CP cottonseed mealbased supplement during grazing of DNR (CON); 2) corn/soybean meal-based supplement fed at 1% of BW during grazing of DNR (CORN); 3) grazing WP at a high stocking rate to achieve a reduced rate of BW gain (LGWP); and 4) grazing WP at a low stocking rate to achieve an increased rate of BW gain (HGWP). In Exp. 1, 3 steers per treatment were harvested after winter grazing (138 d). The remaining WP steers were transitioned into a finishing phase and DNR steers were allowed to graze the same pastures for another 115 d before entering a feedyard. In Exp. 2, steers grazed respective pastures until each treatment reached an estimated HCW of 200 kg (262, 180, 142, and 74 d, respectively, for CON, CORN, LGWP, and HGWP treatments), at which time 4 steers per treatment were randomly selected for intermediate harvest before finishing. At the end of the finishing period, 4 additional steers from each treatment were randomly selected for final carcass measurements. All steers were fed to a common 12th rib fat thickness of 1.27 cm. After winter grazing in Exp. 1, HGWP steers had the greatest (P < 0.01) mesenteric/omental fat, total viscera, total splanchnic tissue mass, and carcass and empty body fat, compared with the other treatments. In Exp. 2 at intermediate harvest, WP steers had greater (P < 0.03) mesenteric/omental fat, total viscera, and total splanchnic tissue mass, compared with CORN steers, with CON steers being intermediate. Also, the WP steers had greater (P < 0.02) carcass and empty body fat, compared with CORN steers, with CON steers being intermediate. At final harvest in Exp. 2, LGWP steers had the least total viscera and total splanchnic tissue mass, compared with the other treatments. However, there were no differences (P > 0.53) among treatments for carcass or empty body fat. Stocker systems using WP or DNR result in cattle with differences in body fat and visceral organ mass before finishing; this may influence feedlot efficiency, even though there were no differences in body fat and visceral organ mass at the end of the finishing period. Key words: adipose deposition, carcass density, growing beef cattle, organ mass 2013 American Society of Animal Science. All rights reserved. J. Anim. Sci : doi: /jas INTRODUCTION Numerous growing programs are used for beef cattle in the Southern Great Plains; these can produce a wide range in ADG that can alter body fat content (i.e., fleshiness) at the start of the finishing phase (Hersom et al., 2004a; Sharman et al., 2013). Fox et 1 Corresponding author: gerald.horn@okstate.edu Received May 8, Accepted February 12, al. (1972) and Oltjen and Garrett (1988) reported that nutritionally restricted steers have increased protein deposition during the first part of finishing, but at the end of finishing, such steers deposit fat at a greater rate compared with steers that have never been restricted nutritionally. Hersom et al. (2004a) reported that steers wintered on wheat pasture at a high rate of gain enter the finishing phase with a greater body fat mass but have similar finishing ADG and similar final marbling scores as compared with steers wintered on dormant native range (DNR).

2 2356 Sharman et al. Hersom et al. (2004a) also demonstrated that the maintenance requirement of growing beef cattle can be influenced by their previous nutritional program. Ferrell and Jenkins (1985) reported that maintenance energy expenditures can be attributed largely to energy use by visceral organs. McLeod and Baldwin (2000) summarized that the gastrointestinal tract (GIT) and liver have considerable impact on ME use and account for 40 to 50% of whole body heat production and varies in regard to ME intake of the diet. Sainz and Bentley (1997) indicated that the amount of absorbed nutrients is the driving factor influencing liver mass, whereas the forestomach and intestine mass respond to dietary fiber content, as well as the amount of absorbed nutrients. Therefore, we hypothesized that steers consuming lower quality forage providing lower ADG (e.g., DNR) would have greater visceral organ mass but lower fat deposition compared with steers consuming greater quality forage providing greater ADG (e.g., winter wheat pasture; WP) during the stocker phase, thereby impacting efficiency during the finishing phase. The objective of the study was to examine the effect of rate of BW gain during the stocker phase on the mass of visceral organs and body composition of steers during the stocker and finishing phases. MATERIALS AND METHODS Before the initiation of these studies, procedures for animal care, handling, and sampling were approved by the Oklahoma State University Institutional Animal Care and Use Committee. Animals, Treatments, and Management Experiments 1 and 2. A detailed description of the stocker and finishing phases has been reported in a companion paper by Sharman et al. (2013) and will be summarized here. Fall-weaned Angus steers from the same Oklahoma State University cow herd were used for 2 separate experiments in consecutive years. In Exp. 1, 72 steers (258 ± 29 kg; 276 ± 18 d of age) and in Exp. 2, 76 steers (258 ± 28 kg; 265 ± 20 d of age) were allotted to graze either DNR or WP. The 4 treatments included: 1) control, 1.02 kg steer 1 d 1 of a 40% CP cottonseed meal-based supplement to meet their degradable intake protein (DIP) requirement during grazing of DNR (CON); 2) ground corn/ soybean meal-based supplement fed at 1% of BW during grazing of DNR (CORN); 3) grazing WP at a high stocking rate (3 steers/ha) to achieve a reduced rate of BW gain (LGWP); and 4) grazing WP at a low stocking rate (1 steer/ha) to achieve an increased rate of BW gain (HGWP). Both experiments were initiated in early December when steers were allotted to treatments based on initial BW and sire information of the steers conceived using AI. At this time, 4 randomly selected steers were harvested at the Oklahoma State University Food and Agricultural Products Center (FAPC) abattoir in Stillwater to collect initial body composition. For each experiment and immediately on hide removal and evisceration, GIT was collected and separated (into the reticulo-rumen, omasum, abomasum, and small and large intestines), emptied of contents, and weighed. Internal organs (heart, lungs, kidney, liver, spleen, and pancreas) and mesenteric/omental fat were also weighed. Empty BW was calculated from a published equation relating HCW with empty BW (Fox et al., 1976), because the hide, head, feet, ears, and blood were not weighed. Total GIT was calculated as the sum of the empty reticulo-rumen, omasum, abomasum, small intestine, and large intestine. Total viscera was calculated as the sum of the total GIT plus mesenteric/omental fat. Total splanchnic tissue was calculated as total viscera plus liver, spleen, and pancreas. After a chill time of 48 h, the left side of each eviscerated carcass was quartered and carcass density measurements were made by the water-displacement method as described by Garrett and Hinman (1969). Each quarter was weighed in air and then submerged in a tank ( cm) containing water at 4 C and weighed, using a triple-beam balance that was placed on a solid platform above the tank. The air temperature in the room was 4 C to eliminate specific water volume adjustments. In Exp. 2, KPH fat was removed from the carcass after evisceration and weighed, placed in a bag, and chilled with the carcasses. Carcass density measurements were as described in Exp. 1, except the density of KPH was measured separately in a wire basket ( cm). The weight of KPH in air and in water was added to the weight of the carcass for calculation of carcass density. Carcass composition for both experiments was calculated from carcass density values, using the equations by Garrett and Hinman (1969). Carcass and empty body fat composition was also calculated from measurements of mesenteric fat, 12th rib fat thickness, and KPH, using equations reported by Sainz et al. (1995; Table 1). Grazing and Finishing Phases In Exp. 1, at the end of the 138-d winter grazing phase, 3 steers were selected from each treatment using herd sire information from AI and calving records, and final grazing BW for the first intermediate harvest. Steers were harvested as described for the initial harvest in Exp. 1. Additionally, the reticulo-ruminal contents were collected and weighed, and 3 subsamples were

3 Organ Mass and Carcass Composition of Stocker Cattle 2357 collected for DM analysis (oven drying at 115 C) to determine the amount of ruminal contents (g/kg of BW) at the end of winter grazing. At this time, the remaining 28 WP steers were transitioned into the finishing phase, whereas the remaining 22 DNR steers were transitioned into a season-long summer grazing program on the same pastures used during winter grazing; winter supplements were not fed. At the end of summer grazing, 3 steers from only CON and CORN treatments were randomly selected, based on BW, and were harvested as previously described. The remaining 8 steers within each of these 2 treatments were transitioned into the finishing phase. All treatments were fed to an estimated 12th rib fat thickness of 1.27 cm. At the end of finishing, 3 steers, selected based on BW from LGWP and HGWP treatments, were harvested, as previously described. No steers from CON and CORN treatments were harvested for collection of visceral organ mass and carcass density measurements, due to scheduling conflicts. In Exp. 2, steers grazed their respective pastures until each treatment reached an estimated HCW of 200 kg. At this time, 4 steers, selected from each treatment using herd sire information from AI and calving records, and final grazing BW, were harvested for intermediate visceral organ and body composition measurements. Steers were harvested as described for the intermediate harvest in Exp. 1. The remaining steers from each treatment were transitioned into the finishing phase. Steers were fed to a predicted common fat endpoint of 1.27 cm of rib fat thickness, based on ultrasound measurements for 12th rib fat thickness that were determined ~1 mo before harvest for assigning harvest date and selection of 4 steers for harvest at the FAPC abattoir. Steers, selected based on their 12th rib fat thickness and final finishing BW, were harvested, as previously described. Statistical Analysis For each experiment, mass of the organs from each harvest was expressed as grams per kilogram of empty BW. Mass of body components and carcass characteristics were analyzed as a completely randomized design with steer as the experimental unit (PROC MIXED; SAS Inst. Inc., Cary, NC). The statistical model included the fixed effect of winter grazing treatment and least squares means were calculated and compared using a protected LSD (P < 0.05). RESULTS Growth performance and carcass data from the 2 experiments have been reported in the companion paper by Sharman et al. (2013). Table 1. Equations used to calculate carcass and empty BW chemical composition using carcass density or carcass characteristics Chemical Component Equation Garrett and Hinman, (1969) Carcass ether extract, % Y = ( carcass density) Carcass water, % Y = ( carcass density) Carcass nitrogen, % Y = (20.00 carcass density) Carcass energy, kcal/gm Y = (43.63 carcass density) Sainz et al. (1995) Carcass fat, 1 % Y = (0.541 mesenteric fat) + ( th rib fat thickness) + (1.58 KPH) Empty body fat, 1 % Y = (0.513 mesenteric fat) + ( th rib fat thickness) + (1.48 KPH) 1 Units for calculation: mesenteric/omental fat = kg; 12th rib fat thickness = mm; KPH = %. Mass of Body Components Initial Harvest. Initial mass of body components for Exp. 1 and 2 are shown in Table 2. Due to using steers from the same cow herd 2 consecutive years and initiating experiments at a similar age of the animal, data are very similar among years. The steers that were selected for initial harvest in Exp. 2 did have more mesenteric/omental fat as a proportion of empty BW and greater 12th rib fat thickness, compared with the initial harvest in Exp. 1 (Sharman et al., 2013). This increase in fat deposition increased the total visceral and total splanchnic tissue mass as a proportion of empty BW, compared with Exp. 1. Intermediate Harvest, Exp. 1. When steers were harvested at a similar age after winter grazing, HGWP steers had greater (P < 0.03) live BW and empty BW, compared with CON and CORN steers, and LGWP steers were intermediate (Table 3). When CON and CORN steers were harvested after summer grazing at the second intermediate harvest, there was no difference in live BW or empty BW, compared with the 2 WP treatments. At the end of winter grazing (first intermediate harvest), DNR steers had greater (P < 0.01) ruminal content (g/kg of BW) compared with WP steers. At first intermediate harvest, carcass mass (g/kg of empty BW) followed a similar trend to empty BW, most likely due to the use of carcass weight to calculate empty BW; HGWP steers had greater (P < 0.01) carcass mass compared with CON and CORN steers, and LGWP steers were intermediate (Table 3). The HGWP steers had greater (P < 0.05) total splanchnic tissue mass (g/kg empty BW) than the other treatments, which is primarily due to differences in total viscera but also pancreas and liver mass. The HGWP steers had greater (P < 0.05) pancreas mass (g/kg empty BW) than CON and LGWP steers, with CORN steers being intermediate. Steers grazing WP had greater (P < 0.05) liver mass (g/kg empty

4 2358 Sharman et al. Table 2. Mass of body components for the initial harvest steers in Exp. 1 and 2 Experiment 1 Experiment 2 Mean SD Mean SD Steer, no. 4 4 Live BW, 1 kg Empty BW, kg g/kg of empty BW Carcass Heart Lungs-trachea Reticulo-rumen Omasum Abomasum Small intestine Large intestine Total GIT Mesenteric/omental fat Total viscera Pancreas Spleen Liver Total splanchnic tissue Kidney Kidney and pelvic fat Live BW = 5-h shrunk live BW with feed and water withheld. 2 Total GIT (gastrointestinal tract) was calculated as the sum of reticulorumen, omasum, abomasum, small intestine, and large intestine. 3 Total viscera was calculated as total GIT plus mesenteric/omental fat. 4 Total splanchnic tissue was calculated as total viscera plus liver, spleen, and pancreas. BW) than steers grazing DNR. At the second intermediate harvest, CON and CORN steers had similar total splanchnic tissue mass, compared with LGWP steers but less (P < 0.05) than HGWP steers. Additionally, CON and CORN steers at the second intermediate harvest had similar kidney mass (g/kg empty BW) compared with CON steers at the first intermediate harvest, which had lower (P < 0.05) kidney mass than the other 3 treatments at the first intermediate harvest. At the first intermediate harvest, HGWP steers had greater (P < 0.05) total viscera (g/kg of empty BW) than the other treatments, which was primarily due to differences in mesenteric/omental fat mass but also omasal and abomasal mass. Mesenteric/omental fat mass (g/kg empty BW) was greatest for HGWP steers, followed by LGWP steers and CON and CORN steers were similar. Mass of the omasum (g/kg of empty BW) was greater (P < 0.05) for CON steers compared with the other treatments. Mass of the abomasum (g/kg empty BW) was greater (P < 0.05) for CORN steers than WP steers, with CON steers being intermediate. Additionally, CON and CORN steers tended (P = 0.08) to have greater large intestinal mass (g/kg of empty BW), compared with the 2 WP treatments. There was a trend (P = 0.06) for CON and HGWP steers to have greater lungs-trachea mass (g/kg of empty BW) compared with CORN and LGWP steers. There were no differences (P > 0.10) among treatments for reticulo-rumen, small intestine, or total GIT mass (g/kg of empty BW). At the second intermediate harvest, when CON and CORN steers had achieved similar (P > 0.10) live BW to LGWP and HGWP steers, respectively, CON and CORN steers had similar (P > 0.10) carcass mass as a proportion of empty BW as LGWP and HGWP steers at first intermediate harvest, respectively. A similar trend was observed for total viscera (g/kg empty BW), where CON and CORN steers had similar (P > 0.10) total visceral mass, compared with LGWP and HGWP steers, respectively. When comparing components of total viscera, CON steers had similar (P > 0.10) mesenteric/omental fat and omasal and abomasal mass as LGWP steers at the first intermediate harvest. However, CORN steers had decreased (P < 0.05) mesenteric/omental fat mass, similar abomasal mass, and greater (P < 0.05) omasal mass (g/kg empty BW), compared with HGWP steers at the first intermediate harvest. Intermediate Harvest, Exp. 2. When steers were harvested at a similar HCW (200 kg), there were no differences (P > 0.52) in live or empty BW (Table 4). The CON and CORN steers had greater (P < 0.03) ruminal content (g/kg of BW), compared with LGWP steers, with HGWP steers being intermediate, which agrees with the results observed in Exp. 1 at the end of winter grazing. There were no differences in carcass mass (g/kg empty BW) among treatments. Total splanchnic tissue mass (g/kg empty BW) was less (P < 0.05) for CORN steers than CON and HGWP steers, with LGWP steers being intermediate. There were no differences in mass of pancreas, spleen, and liver, indicating that differences in total splanchnic tissue mass were due to differences in visceral components. Even though there were no differences in liver mass, the trend for treatment means is similar to Exp. 1, where WP steers had greater liver mass than DNR steers. Total viscera (g/kg empty BW) was less (P < 0.05) for CORN steers compared with CON and HGWP steers; LGWP steers were intermediate, which may be due to differences in mesenteric/ omental fat and omasal mass. The CORN steers tended (P < 0.07) to have less mesenteric/omental fat mass (g/kg empty BW) compared with the other treatments. The CORN steers had decreased (P < 0.05) omasal mass (g/kg empty BW) than CON steers but greater omasal mass than HGWP steers. There were no differences in kidney mass, but LGWP and HGWP steers had greater (P < 0.05) kidney and pelvic fat mass (g/kg empty BW) than CORN steers, with CON being intermediate. Final Harvest, Exp. 1. Only LGWP and HGWP treatments were harvested at the FAPC abattoir to

5 Organ Mass and Carcass Composition of Stocker Cattle 2359 Table 3. Mass of body components for steers from different stocker production programs at the first and second intermediate harvest in Exp. 1 First intermediate 2 Second intermediate 2 CON CORN LGWP HGWP CON CORN SEM P-value Steers, no Live BW, 3 kg 286 a 324 ab 354 abc 401 c 362 bc 405 c Ruminal content, g/kg BW 9.6 b 8.3 b 4.5 a 5.4 a Empty BW, kg 251 a 292 ab 334 bc 373 c 303 b 332 bc g/kg of empty BW Carcass 599 a 615 ab 628 bc 637 c 619 b 627 bc Heart 6.2 a 6.5 a 8.6 b 7.7 ab 5.9 a 5.9 a Lungs-trachea Reticulo-rumen Omasum 10.7 b 8.3 a 7.9 a 6.9 a 10.1 b 10.1 b Abomasum 4.9 bc 5.3 c 4.1 ab 4.2 ab 3.3 a 3.3 a Small intestine Large intestine Total GIT Mesenteric/omental fat 6.2 a 8.9 a 15.9 b 29.1 c 15.7 b 18.1 b Total viscera ab 72.3 a 74.7 ab 86.7 c 76.9 ab 80.1 bc Pancreas 0.9 a 1.2 ab 1.0 a 1.5 b 1.0 a 1.3 b Spleen Liver 14.1 a 13.0 a 16.7 b 16.6 b 13.3 a 13.4 a Total splanchnic tissue ab 88.4 a 95.0 ab c 93.4 ab 96.9 b Kidney 3.2 ab 3.7 b 4.8 c 5.1 c 2.5 a 2.3 a CON = 40% CP supplement; CORN = corn-based supplement; LGWP = reduced rate of BW gain on wheat pasture; HGWP = increased rate of BW gain on wheat pasture. 2 First intermediate harvest was conducted at the end of winter grazing; second intermediate harvest was conducted at the end of summer grazing. 3 Live BW = 5-h shrunk live BW with feed and water withheld. 4 Total GIT (gastrointestinal tract) was calculated as the sum of reticulo-rumen, omasum, abomasum, small intestine, and large intestine. 5 Total viscera was calculated as total GIT plus mesenteric/omental fat. 6 Total splanchnic tissue was calculated as total viscera plus liver, spleen, and pancreas. a c Within a row, means without a common superscript differ (P < 0.05). collect mass of body components, due to scheduling conflicts for CON and CORN treatments (Table 5). There was no difference (P > 0.87) between treatments for live BW and empty BW. The HGWP steers had greater (P < 0.05) total splanchnic tissue and total visceral mass (g/kg empty BW), primarily due to greater (P < 0.05) mesenteric/omental fat mass (g/kg empty BW), but also the tendency (P = 0.10) for greater reticulo-rumen and small intestinal mass (g/kg empty BW), compared with LGWP steers. The HGWP steers also tended (P = 0.07) to have greater kidney mass (g/ kg empty BW) than LGWP steers. There were no differences (P > 0.10) in pancreas, spleen, or liver mass among treatments. Also, there was no difference (P > 0.32) in carcass, heart, or lungs-trachea mass (g/kg of empty BW) between the WP treatments. Final Harvest, Exp. 2. There were no differences (P > 0.10) among the treatments for live BW, empty BW, or carcass weight at final harvest (Table 6). The CON steers had greater (P < 0.01) total splanchnic tissue mass (g/kg of empty BW) compared with LGWP steers, with CORN and HGWP steers being intermediate, which is primarily due to differences in visceral components but also spleen and liver mass. The CON steers had greater (P < 0.05) spleen and liver mass (g/kg empty BW), compared with the other treatments. The CORN and LGWP steers had less (P < 0.05) total visceral mass (g/kg of empty BW), compared with CON steers; HGWP steers were intermediate, which is due to differences in reticulo-rumen and omasal mass, and mesenteric/omental fat mass. The HGWP steers had greater (P < 0.01) reticulo-rumen mass (g/kg of empty BW) compared with LGWP steers, with CON and CORN steers being intermediate, which is similar to the differences between HGWP and LGWP steers observed in Exp. 1. The WP steers had less (P < 0.01) omasal mass (g/kg of empty BW) compared with the DNR steers. These differences resulted in a trend (P = 0.09) for LGWP steers to have lower total gastrointestinal tract mass (g/kg empty BW) than the other treatments. The LGWP steers also tended (P = 0.08) to have lower mesenteric/omental fat mass (g/kg empty BW) than CON and HGWP steers. There

6 2360 Sharman et al. Table 4. Mass of body components for steers from different stocker production programs at the intermediate harvest in Exp. 2 SEM P-value CON CORN LGWP HGWP Steers, no Live BW, 2 kg Ruminal content, g/kg BW 9.2 b 8.6 b 5.4 a 7.5 ab Empty BW, kg g/kg of empty BW Carcass Heart 5.9 b 5.8 b 4.8 a 6.2 b Lungs-trachea Reticulo-rumen Omasum 12.0 c 8.8 b 7.2 ab 6.9 a Abomasum Small intestine Large intestine Total GIT Mesenteric/omental fat Total viscera b 75.0 a 77.4 ab 84.9 b Pancreas Spleen Liver Total splanchnic tissue b 92.4 a 95.9 ab b Kidney Kidney and pelvic fat 5.7 ab 4.4 a 8.2 c 7.4 bc CON = 40% CP supplement; CORN = corn-based supplement; LGWP = reduced rate of BW gain on wheat pasture; HGWP = increased rate of BW gain on wheat pasture. 2 Live BW = 5-h shrunk live BW with feed and water withheld. 3 Total GIT (gastrointestinal tract) was calculated as the sum of reticulorumen, omasum, abomasum, small intestine, and large intestine. 4 Total viscera was calculated as total GIT plus mesenteric/omental fat. 5 Total splanchnic tissue was calculated as total viscera plus liver, spleen, and pancreas. a c Within a row, means without a common superscript differ (P < 0.05). were no differences (P > 0.13) among the treatments for pancreas, kidney, or kidney and pelvic fat mass (g/kg of empty BW) at final harvest. There was also no difference (P > 0.27) for heart or lungs-trachea mass (g/kg of empty BW) among treatments. Carcass and Empty Body Weight Chemical Composition Initial Harvest. The carcass and empty BW chemical composition for steers at the initial harvests for Exp. 1 and 2 is shown in Table 7. The steers in Exp. 2 had considerably more 12th rib fat thickness compared with Exp. 1. Additionally, the initial harvest steers in Exp. 2 had much lower carcass density values, which led to greater carcass fat, based on the equations by Garrett and Hinman (1969) for chemical composition. The values derived using the Garrett and Hinman (1969) equations are much greater Table 5. Mass of body components for steers from different wheat pasture stocker programs at final harvest in Exp. 1 LGWP HGWP SEM P-value Steers, no. 3 3 Live BW, 2 kg Empty BW, kg g/kg of empty BW Carcass Heart Lungs-trachea Reticulo-rumen Omasum Abomasum Small intestine Large intestine Total GIT Mesenteric/omental fat Total viscera Pancreas Spleen Liver Total splanchnic tissue Kidney LGWP = reduced rate of BW gain on wheat pasture; HGWP = increased rate of BW gain on wheat pasture. 2 Live BW = 5-h shrunk live BW with feed and water withheld. 3 Total GIT (gastrointestinal tract) was calculated as the sum of reticulorumen, omasum, abomasum, small intestine, and large intestine. 4 Total viscera was calculated as total GIT plus mesenteric/omental fat. 5 Total splanchnic tissue was calculated as total viscera plus liver, spleen, and pancreas. than values derived using equations by Sainz et al. (1995). The values derived using equations published by Sainz et al. (1995) are similar to data reported by Sainz et al. (1995) for weaned steers (BW = 245 kg) but less than that reported by McCurdy et al. (2010b). Intermediate Harvest, Exp. 1. As previously reported by Sharman et al. (2013), HGWP steers had greater (P < 0.01) HCW, 12th rib fat thickness, KPH, mesenteric/ omental fat, and marbling score, compared with the other grazing treatments at first intermediate harvest (Table 8). The LGWP steers had greater HCW, mesenteric/omental fat, and marbling score, compared with CON steers, with CORN steers being intermediate. At the second intermediate harvest, CON and CORN steers had similar HCW, 12th rib fat thickness, and marbling score as CORN and LGWP steers at the first intermediate harvest, suggesting that as steers grew to similar BW on summer pasture, accretion of individual fat depots compensated. There were no differences (P > 0.61) among the treatments at either intermediate harvest for carcass density or for chemical composition components, when derived from the Garrett and Hinman (1969) equations. Based on Sainz et al. (1995) equations, HGWP steers had greater

7 Organ Mass and Carcass Composition of Stocker Cattle 2361 Table 6. Mass of body components for steers from different stocker production programs at final harvest in Exp. 2 SEM P-value CON CORN LGWP HGWP Steers, no Live BW, 2 kg Empty BW, kg g/kg of empty BW Carcass Heart Lungs-trachea Reticulo-rumen 25.0 ab 22.4 ab 21.1 a 25.0 b Omasum 7.1 b 6.4 b 5.5 a 5.2 a Abomasum Small intestine Large intestine Total GIT Mesenteric/omental fat Total viscera b 90.0 a 84.4 a 96.7 ab Pancreas Spleen 2.2 b 1.7 a 1.8 a 1.6 a Liver 17.5 b 14.0 a 14.7 a 15.0 a Total splanchnic tissue c ab a bc Kidney Kidney and pelvic fat CON = 40% CP supplement; CORN = corn-based supplement; LGWP = reduced rate of BW gain on wheat pasture; HGWP = increased rate of BW gain on wheat pasture. 2 Live BW = 5-h shrunk live BW with feed and water withheld. 3 Total GIT (gastrointestinal tract) was calculated as the sum of reticulorumen, omasum, abomasum, small intestine, and large intestine. 4 Total viscera was calculated as total GIT plus mesenteric/omental fat. 5 Total splanchnic tissue was calculated as total viscera plus liver, spleen, and pancreas. a c Within a row, means without a common superscript differ (P < 0.05). (P < 0.02) carcass and empty body fat, compared with the other treatments at both intermediate harvests. The CON and CORN steers at the first intermediate harvest had the lowest carcass and empty body fat, but at the second intermediate harvest (end of summer grazing), there were no differences in body fat content compared with LGWP and CORN steers at first intermediate harvest. Intermediate Harvest, Exp. 2. Steers were similar in HCW (Table 9), except for LGWP steers that tended (P = 0.09) to be slightly heavier at intermediate harvest due to an unexpected increase in rate of gain the last 2 wk of the grazing phase caused by rapid forage growth. There was a tendency (P = 0.06) for HGWP steers to have greater 12th rib fat thickness when compared with CORN steers, with CON and LGWP being intermediate. The WP steers had greater (P < 0.01) KPH but were similar (P > 0.10) in greater mesenteric/omental fat, compared with DNR steers. The LGWP steers had greater (P < 0.01) marbling score compared with HGWP steers and HGWP steers had greater marbling score than CON and CORN steers. Table 7. Carcass and empty BW chemical composition for the initial harvest steers in Exp. 1 and 2 Experiment 1 Experiment 2 Mean SD Mean SD Steer, no. 4 4 HCW, kg th rib fat thickness, mm KPH, % Mesenteric/omental fat, kg Marbling score Carcass density Garrett and Hinman, (1969) Carcass ether extract, % Carcass water, % Carcass nitrogen, % Carcass energy, kcal/gm Sainz et al. (1995) Carcass fat, % Empty body fat, % Marbling score grid: 100 = practically devoid 00 ; 200 = traces 00 ; 300 = slight 00. When values were derived from Garrett and Hinman (1969) equations, there were no differences (P > 0.21) in carcass density or chemical composition among treatments when harvested at a similar HCW before entering the finishing phase. The HGWP steers had a numerically greater percentage of fat and energy content with lower water and nitrogen content, compared with CON, CORN, and LGWP steers, which were similar. When using Sainz et al. (1995) equations, WP steers had greater (P < 0.02) carcass and empty body fat percentage, compared with CORN steers, with CON steers being intermediate. Final Harvest, Exp. 1. Final carcass characteristics are reported by Sharman et al. (2013), which includes steers harvested at the FAPC abattoir and a commercial abattoir. Data reported here represents only those steers harvested at FAPC that were used for determination of carcass density measurements (Table 10). The HGWP steers tended (P = 0.10) to have greater KPH compared with the LGWP steers. However, there were no differences in carcass and empty BW chemical composition between the 2 WP treatments at the end of finishing. Final Harvest, Exp. 2. Final carcass characteristics are reported by Sharman et al. (2013), which includes steers harvested at the FAPC abattoir and a commercial abattoir. Data reported here represent only those steers harvested at the FAPC abattoir that were used for determination of carcass density measurements (Table 11). The CON steers tended (P = 0.09) to have lower 12th rib fat thickness compared with the other treatments. However, there were no differences in HCW, KPH, or mesenteric/omental fat among treatments. The HGWP steers that were harvested at the FAPC abattoir had lower (P < 0.04) marbling score compared with CON and CORN steers, with LGWP steers being intermediate.

8 2362 Sharman et al. Table 8. Carcass and empty BW chemical composition for steers from different stocker production programs at the first and second intermediate harvest in Exp. 1 First intermediate 2 Second intermediate 2 CON CORN LGWP HGWP CON CORN SEM P-value Steers, no HCW, kg 150 a 180 ab 210 b 238 c 188 b 208 bc th rib fat thickness, mm 0.34 a 1.02 ab 1.69 ab 8.47 c 1.69 ab 2.71 b KPH, % 0.50 a 0.50 a 0.67 a 1.33 b 0.67 a 0.83 a Mesenteric/omental fat, kg 1.53 a 2.64 a 5.34 b c 4.74 b 5.99 b Marbling score a 217 ab 280 c 340 d 233 ab 240 bc Carcass density Garrett and Hinman, (1969) Carcass ether extract, % Carcass water, % Carcass nitrogen, % Carcass energy, kcal/gm Sainz et al. (1995) Carcass fat, % 10.5 a 11.4 ab 13.5 bc 21.0 d 13.2 bc 14.6 c Empty body fat, % 8.8 a 9.7 ab 11.6 c 18.6 d 11.3 bc 12.7 c CON = 40% CP supplement; CORN = corn-based supplement; LGWP = reduced rate of BW gain on wheat pasture; HGWP = increased rate of BW gain on wheat pasture. 2 First intermediate harvest was conducted at the end of winter grazing; second intermediate harvest was conducted at the end of summer grazing. 3 Marbling score grid: 100 = practically devoid 00 ; 200 = traces 00 ; 300 = slight 00 ; 400 = small 00. a d Within a row, means without a common superscript differ (P < 0.05). The LGWP steers tended (P = 0.06) to have larger carcass density values, which resulted in reduced carcass fat and energy content, and greater water and nitrogen content. However, there were no differences (P > 0.53) in carcass or empty body fat content among treatments using equations developed by Sainz et al. (1995). DISCUSSION Growth performance and carcass characteristics for Exp. 1 and 2 are presented by Sharman et al. (2013). Only data pertaining to steers harvested at the FAPC abattoir will be discussed in this manuscript. Mass of Body Components Hersom et al. (2004b) reported that when steers were harvested at a similar age just before finishing, steers previously grazing WP at an increased rate of BW gain had greater empty BW compared with steers grazing DNR, which is similar to the first intermediate harvest in Exp. 1. However, at final harvest, there were no differences in empty BW among the treatments, which is similar to the current experiments. McCurdy et al. (2010b) also reported no difference in empty BW percentage or change in empty BW from the end of the growing phase at intermediate harvest to the end of finishing among steers grazing WP, fed a silage-based diet ad libitum, or programmed fed a concentrate diet during the growing phase. In Exp. 2, empty BW was similar among the treatments at both the intermediate and final harvests, which demonstrates a similar change in empty BW as shown by McCurdy et al. (2010b). In the current experiments, WP steers had greater total splanchnic tissue mass compared with steers grazing DNR at both first and second intermediate harvests in Exp. 1, which was due to greater liver mass of WP steers compared with DNR steers and the differences in total viscera previously mentioned. In Exp. 2, WP steers had greater liver mass compared with DNR steers; however, only CORN steers had lower total splanchnic tissue mass compared with WP steers at intermediate harvest. At final harvest in Exp. 2, CON steers had greater liver mass than the other treatments and greater total splanchnic tissue mass, compared with CORN and LGWP steers. Hersom et al. (2004b) reported that nutritionally restricted steers had lower liver mass compared with steers grazing WP to produce a high rate of BW gain in yr 1, but no differences were observed in yr 2. McCurdy et al. (2010b) observed no difference in liver and total splanchnic tissue mass among growing programs managed for similar rates of gain at the end of the growing or finishing phases. These results agree with the concept that liver mass responds to increases in total energy intake and metabolic demand (Sainz and Bentley, 1997; Hersom et al., 2004b), but does not respond to type of diet that was fed previously (McLeod and Baldwin, 2000).

9 Organ Mass and Carcass Composition of Stocker Cattle 2363 Table 9. Carcass and empty BW chemical composition for steers from different stocker production programs at intermediate harvest in Exp. 2 Table 10. Carcass and empty BW chemical composition for steers from wheat pasture stocker programs at final harvest in Exp. 1 SEM P-value CON CORN LGWP HGWP Steers, no HCW, kg th rib fat thickness, mm KPH, % 0.63 a 0.50 a 1.13 b 1.38 b Mesenteric/omental fat, kg Marbling score a 143 a 315 c 228 b Carcass density Garrett and Hinman, (1969) Carcass ether extract, % Carcass water, % Carcass nitrogen, % Carcass energy, kcal/gm Sainz et al. (1995) Carcass fat, % 15.0 ab 12.3 a 16.3 b 17.3 b Empty body fat, % 13.0 ab 10.5 a 14.2 b 15.2 b CON = 40% CP supplement; CORN = corn-based supplement; LGWP = reduced rate of BW gain on wheat pasture; HGWP = increased rate of BW gain on wheat pasture. 2 Marbling score grid: 100 = practically devoid 00 ; 200 = traces 00 ; 300 = slight 00 ; 400 = small 00. a c Within a row, means without a common superscript differ (P < 0.05). SEM P-value LGWP HGWP Steers, no. 3 3 HCW, kg th rib fat thickness, mm KPH, % Mesenteric/omental fat, kg Marbling score Carcass density Garrett and Hinman, (1969) Carcass ether extract, % Carcass water, % Carcass nitrogen, % Carcass energy, kcal/gm Sainz et al. (1995) Carcass fat, % Empty body fat, % LGWP = reduced rate of BW gain on wheat pasture; HGWP = increased rate of BW gain on wheat pasture. 2 Marbling score grid: 300 = slight 00 ; 400 = small 00. In the current experiments, differences among treatment were observed for total visceral mass, primarily due to differences in mesenteric/omental fat mass; but components of GIT also contributed to those differences. Total visceral mass was greatest for HGWP steers in Exp. 1 compared with the other treatments at first and second intermediate harvests. In Exp. 2, CORN steers had lower total visceral mass at intermediate harvest compared with CON and HGWP steers, when harvested at a similar HCW. In contrast, McCurdy et al. (2010b) reported no differences in total visceral mass before finishing or at the end of finishing among different growing programs managed for similar rates of BW gain. At the first intermediate harvest in Exp. 1, HGWP steers had greater mesenteric/omental fat compared with LGWP steers, and LGWP steers had greater mesenteric/ omental fat compared with DNR steers. However, CON and CORN steers at the second intermediate harvest had similar mesenteric/omental fat, compared with LGWP steers before entering the finishing phase. In Exp. 2, CORN steers tended to have reduced mesenteric/omental fat compared with LGWP and HGWP steers, when harvested at a similar HCW before entering the finishing phase, which could be due to the steers not having enough days during summer grazing to compensate for the nutritional restriction during winter grazing. Similarly, Hersom et al. (2004b) reported that at the end of winter grazing, steers grazing WP to produce an increased rate of BW gain had greater mesenteric/omental fat mass compared with steers grazing WP to produce a low rate of BW gain, with steers grazing DNR having the least amount of mesenteric/omental fat. McCurdy et al. (2010b) observed that steers programmed fed a highconcentrate diet had greater mesenteric/omental fat at the end of the growing phase compared with steers that grazed WP. Sainz et al. (1995) reported that mesenteric/ omental fat increased as empty BW increased in steers either grown on a high-concentrate diet fed ad libitum, programmed fed a high-concentrate diet, or fed a foragebased diet ad libitum. In both Exp. 1 and 2, HGWP steers had greater mesenteric/omental fat than LGWP steers at final harvest. In contrast to the current experiments, Hersom et al. (2004b) and McCurdy et al. (2010b) observed no difference at the end of finishing for mesenteric/omental fat deposition among different growing programs managed for similar rates of BW gain. Hersom et al. (2004b) reported that steers grazing DNR and WP to produce a low rate of BW gain had increased reticulo-rumen mass at intermediate harvest compared with steers grazing WP to produce a high rate of BW gain. These data suggest that this was caused by reduced energy intake corresponding to greater intake of low-quality forage (Hersom et al., 2004b). Similarly, Sainz and Bentley (1997) showed that steers fed a forage-based diet during the growing phase had greater forestomach mass compared with steers grown on a concentrate diet. However, in the current experiments, there were no differences among grazing treatments for reticulo-rumen mass (g/kg of empty BW) at intermediate

10 2364 Sharman et al. Table 11. Carcass and empty BW chemical composition for steers from different stocker production programs at final harvest in Exp. 2 SEM P-value CON CORN LGWP HGWP Steers, no HCW, kg th rib fat thickness, mm KPH, % Mesenteric/omental fat, kg Marbling score bc 418 c 363 ab 340 a Carcass density Garrett and Hinman, (1969) Carcass ether extract, % Carcass water, % Carcass nitrogen, % Carcass energy, kcal/gm Sainz et al. (1995) Carcass fat, % Empty body fat, % CON = 40% CP supplement; CORN = corn-based supplement; LGWP = reduced rate of BW gain on wheat pasture; HGWP = increased rate of BW gain on wheat pasture. 2 Marbling score grid: 300 = slight 00 ; 400 = small 00 ; 500 = modest 00. a c Within a row, means without a common superscript differ (P < 0.05). harvest. The current experiments weighed the contents (g/kg of BW) within the reticulo-rumen and observed that WP steers had less ruminal content when harvested at a similar age and similar HCW, compared with DNR steers. Even though we saw no differences in reticulorumen mass, contents of the reticulo-rumen correlate with results from Sainz and Bentley (1997) and Hersom et al. (2004b), when steers consuming a greater foragebased diet had greater reticulo-rumen or forestomach mass, respectively. Steers at the first intermediate harvest in Exp. 1 and intermediate harvest in Exp. 2 were harvested with a full GIT, so the contents within the reticulo-rumen provide data indicating that DNR steers had a greater ruminal capacity compared with WP steers, due to consuming a reduced energy diet. Hersom et al. (2004b) reported that DNR steers had heavier omasal mass at entry into the feedlot compared with steers grazing winter WP, which is similar to the current experiments, where omasum mass was greater for CON steers compared with the other treatments. McCurdy et al. (2010b) observed no differences in omasal mass at the end of the growing phase when steers had similar rates of BW gain, even though fiber content of the diet differed (NDF = 45%, 43%, and 21% for WP, silage-based diet, and programmed fed high-concentrate diet, respectively). McLeod and Baldwin (2000) reported that steers fed a forage-based diet had greater omasal and reticular mass (% of BW), compared with steers receiving a concentrate diet. Hersom et al. (2004b) reported no difference in small intestinal mass (g/kg of empty BW) at the end of winter grazing for yr 1; however, in yr 2, researchers observed that nutritionally restricted steers had greater small intestinal mass (g/kg of empty BW) compared with steers grazing WP at a high rate of BW gain. The current experiments are similar to yr 1 of Hersom et al. (2004b) in that no difference was detected in small intestinal mass among any of the treatments at any of the harvest dates. In contrast, Drouillard et al. (1991) observed that intestinal mass of sheep that were nutritionally restricted for 14 or 42 d was less compared with those of unrestricted sheep. McCurdy et al. (2010b) reported that steers in a confinement setting fed a silage-based diet ad libitum or programmed fed a concentrate diet had reduced small intestinal mass at the end of the growing phase compared with steers grazing WP. There were no differences in small intestinal mass at the end of finishing (McCurdy et al., 2010b). Sainz and Bentley (1997) reported that steers previously restricted nutritionally and then alimented to a high-concentrate finishing diet had greater small intestinal mass at the end of finishing compared with steers that were never nutritionally restricted. Hersom et al. (2004b) reported that steers restricted nutritionally had greater total GIT mass (g/kg of empty BW) compared with steers grazing WP to produce a high rate of gain. But, at the end of finishing, there were no differences among treatments for total GIT mass (Hersom et al., 2004b). In the current experiments, total GIT mass was not different among treatments at the first intermediate harvest in Exp. 1 or intermediate harvest in Exp. 2. At the end of finishing in Exp. 2, LGWP steers tended to have a lower total GIT mass than the other treatments but not in Exp. 1. The GIT and liver account for ~10 to 13% of total body mass (Seal and Reynolds, 1993), but account for 40% of the oxygen consumption of the whole body (McBride and Kelly, 1990). Protein synthesis in GIT accounts for ~20 to 23% of the total energy expenditure of GIT (McBride and Kelly, 1990). However, studies have shown that metabolic rate per unit of mass does not change; rather, it is organ mass that changes in response to physiological demand. McBride and Kelly (1990) indicated that, on average, GIT accounts for 7% of empty BW and 20% of whole-body energy expenditure, and the liver accounts for 2% of empty BW and 20% of whole-body energy expenditure. Using these relationships, we estimated that GIT of CON, CORN, LGWP, and HGWP steers at the first intermediate harvest in Exp. 1 would account for 19%, 18%, 17%, and 16% of whole-body energy expenditure, respectively, and that liver would account for 14%, 13%, 17%, and

11 Organ Mass and Carcass Composition of Stocker Cattle % of whole-body energy expenditure, respectively. At the intermediate harvest in Exp. 2, we estimated that the GIT of CON, CORN, LGWP, and HGWP steers would account for 19%, 17%, 16%, and 18% of wholebody energy expenditure, respectively, and that liver would account for 14%, 14%, 15%, and 16% of wholebody energy expenditure, respectively. These differences in organ mass can affect maintenance requirements and could possibly influence efficiency during finishing. In the current experiments, Sharman et al. (2013) reported a tendency for CON steers to have greater G:F during finishing compared with the other treatments in Exp. 1. This corresponds to the reduced liver mass of CON steers at the second intermediate harvest before finishing in Exp. 1, compared with WP steers, which suggests that their maintenance energy requirement may have been lower and they may have been more efficient during finishing. Carcass and Empty Body Weight Chemical Composition Equations relating carcass density (Garrett and Hinman, 1969) and carcass fat characteristics (Sainz et al., 1995) to carcass and empty body chemical composition were used to compute carcass chemical composition. Hersom et al. (2004a) used grazing treatments very similar to those used in Exp. 1 (i.e., HGWP, LGWP, and CON harvested at similar age after winter grazing) and McCurdy et al. (2010a) reported carcass chemical composition of weaned steers and steers after grazing WP to produce an increased rate of BW gain similar to HGWP in Exp. 1. In these studies, chemical composition was determined by proximate analysis of the ground carcass. Hersom et al. (2004a) and McCurdy et al. (2010a) reported that steers grazing WP to produce an increased rate of BW gain had carcass fat content of 20.7% and 15.1%, respectively, which is considerably less than that predicted by the equations of Garrett and Hinman (1969) for HGWP steers at the intermediate harvest with a value of 27.58%. In contrast, equations developed by Sainz et al. (1995) gave a value of 20.97%, which is very similar to that reported by Hersom et al. (2004a), using proximate analysis of the ground carcass. Hersom et al. (2004a) reported carcass fat values of 14.3% and 12.8%, and 7.4% and 4.7% for steers grazing WP to produce a reduced rate of BW gain and steers grazing DNR, respectively, for 2 separate years. In Exp. 1, the equations of Garrett and Hinman (1969) predicted 21.69% and 21.54% carcass fat for LGWP and CON steers, respectively, at the first intermediate harvest; these are considerably greater than the values reported by Hersom et al. (2004a), using proximate analysis of ground carcasses. In contrast, the equations reported by Sainz et al. (1995) predicted carcass fat of 13.48% and 10.47% for LGWP and CON steers at first intermediate harvest. Interestingly, Gil et al. (1970) determined carcass density on 59 carcasses, ranging from 10 to 42% fat and observed that carcass specific volume had a very poor relationship (R 2 = 0.03 to 0.25) with ether extract in lean carcasses (10 to 29% ether extract). Similarly, Carstens et al. (1991) reported that using carcass indicator cuts or carcass density to determine empty BW components may be misleading due to the greater magnitude of composition change observed in the noncarcass tissue components. When evaluating carcass chemical composition at final harvest, using both the equations of Garrett and Hinman (1969) and those of Sainz et al. (1995), it appeared that equations of Sainz et al. (1995) gave results similar to the values reported by Hersom et al. (2004a). Therefore, the equations developed by Sainz et al. (1995), which estimate carcass fat from 10 to 21% for intermediate harvest and from 28 to 34% for final harvest, provide values more similar to the chemical composition values reported by Hersom et al. (2004a) and McCurdy et al. (2010a) than values predicted using the equations of Garrett and Hinman (1969). Hersom et al. (2004a) reported that steers grazing WP to produce an increased rate of BW gain had greater carcass and empty body fat at the end of grazing compared with steers grazing WP to produce a reduced rate of BW gain, with steers grazing DNR having the lowest carcass and empty body fat (averaging 20.14%, 13.33%, 5.80%, respectively). Similarly, in Exp. 1, HGWP steers had the greatest amount of carcass fat at both intermediate harvests compared with the other 3 treatments; however, in Exp. 2, there was no difference between HGWP, LGWP, and CORN steers for carcass or empty body fat when harvested at similar HCW at the end of grazing. Hersom et al. (2004a) reported no differences for carcass or empty body fat composition at the end of finishing in yr 1, but in yr 2, the restricted steers had greater carcass and empty body fat, compared with steers grazing WP to produce an increased rate of BW gain, suggesting a compensatory gain effect. In contrast, Carstens et al. (1991) reported that steers exhibiting compensatory BW gain had ~18% less NEg requirements (kcal/kg EBW) during finishing. At the end of finishing, compensatory steers had less empty body fat and more empty body protein, compared with steers that were continuously grown. Similarly, Drouillard et al. (1991) observed that nutritionally restricted lambs had less empty body fat and greater empty body protein, compared with lambs that were not restricted. Fox et al. (1972) reported that steers exhibiting compensatory gain and grown to 341 kg had greater empty body protein and less empty