Effects of nutrition and management during the stocker phase on quality grade White Paper Certified Angus Beef Clint R. Krehbiel*, Phillip A. Lancaster*, Evin D. Sharman*, Gerald W. Horn*, D. L. Step**, and Deb L. VanOverbeke* *Department of Animal Sciences, Division of Agricultural Sciences and Natural Resources; and **Department of Veterinary Clinical Sciences, Center for Veterinary Health Sciences, Oklahoma State University, Stillwater, 74078 INTRODUCTION The 2011 National Beef Quality Audit Survey reported that “Eating Satisfaction” (defined by packers as tenderness, flavor-marbling) is the second most important attribute in beef consumption behind food safety (Igo et al., 2012). Retained ownership, alliances, and vertically coordinated supply chains are becoming an increasingly larger percentage of cattle on feed, resulting in an estimated 50% or greater of the cattle in the U.S. trading outside of the cash market (Ritchie, 2002). As segments of the beef industry become more coordinated, it becomes increasingly important to understand the effects that management and nutrition in each segment has on all subsequent phases of production, final carcass value, and “Eating Satisfaction” of the final products. Improving nutritional and management strategies for growing beef cattle to enhance final carcass quality will not only benefit the beef industry as a whole, but will provide producers with more incentive to produce high-quality beef to meet consumer demand. Because consumer dollars ultimately drive the beef cattle industry, meeting consumer demand/desires will continue to determine profitability in the future. Studies of nutrition and management practices that influence marbling (intramuscular fat) deposition have primarily focused on the feedlot phase of production (Owens and Gardner, 1 2000). However, pre-feedyard management strategies (health status, stocker/backgrounding, nutrient supplementation, etc.) can influence marbling development (Anderson and Gleghorn, 2007). Therefore, changes in management practices during early phases of the production cycle that increase intramuscular fat deposition and decrease fat deposition in other depots could enhance the efficiency of beef production and enhance carcass quality. Due to the current price of grains and harvested forages, cattle are entering the feedlot at heavier weights indicating that post-weaning management programs are being used to decrease the number of days cattle spend in the feedyard. Understanding how these post-weaning nutrition and management programs impact carcass growth and development is becoming increasingly important. Depending on biological type of cattle, calves are often grazed or placed on a growing diet after weaning to achieve adequate frame size and carcass weight before entering the feedlot for finishing. Across the Southern Great Plains and the Southeastern U.S., grazing systems are commonly used for growing programs during the winter. Cool season forages, including wheat pasture, are utilized to grow cattle to desirable weights for feedlot entry (Byers, 1982). In addition, each year in the Southern Great Plains fall-weaned calves are wintered on dormant native range. Cattle that are wintered on dormant native range are typically fed a protein supplement to gain 0.25 to 0.50 kg/day through the winter months until spring forage growth occurs. These calves then graze summer pasture in either intensive-early stocking or season-long grazing programs prior to entering the finishing phase. In the Northern Great Plains, cattle are usually either grown on crop residues, dormant grass pastures, or placed in confinement and program fed for a moderate rate of gain on harvested and ensiled crops such as corn silage. Albeit stocker programs are increasing due to costs of gain in the feedyard, some larger-framed calves may go directly on to a high-concentrate diet immediately following weaning. High- 2 concentrate diets may be fed at restricted levels in the growing phase to provide a desired rate of gain while allowing for lean tissue growth (Sip and Pritchard, 1991). Because of the variability in growing programs among different regions of the country and in diets that may consist of grazed or harvested forage and/or grain-based diets, performance and weight gain of cattle before and after feedlot entry may be vastly different (McCurdy et al., 2010). In addition, development of fat depots in relation to BW and maturity of the animal could be impacted to alter carcass quality. Because economic value of carcasses is dependent primarily on carcass weight and carcass quality, cattle producers have a strong interest in factors associated with maximizing these two variables. This review will focus on stocker cattle nutrition and management strategies that have the potential to impact marbling score and carcass quality. MARBLING DEPOSITION DURING THE STOCKER PHASE Adipose (fat) tissue development is of major importance to beef production because it influences production efficiency, product quality and consumer acceptance, and, therefore, product value (Smith et al., 1987). Growth of the intramuscular fat depot (i.e., marbling) is especially desirable due to consumer preference for well-marbled beef, whereas growth of other fat depots results in excess fat and production inefficiencies (Hausman et al., 2009). In addition, the amount of fat deposited in the intramuscular depot is the basis for carcass price premiums (marbling; USDA Quality Grade) and fat deposited in other depots can result in carcass discounts (subcutaneous fat; USDA Yield Grade). Anderson and Gleghorn (2007) suggested that marbling deposition is a lifetime event and that pre-feedlot nutrition has a significant impact on marbling deposition. Peel (2003) estimated that 76% of the yearly calf crop enters a backgrounding or stocker program prior to finishing; therefore, there is tremendous opportunity to improve carcass quality attributes by influencing adipose tissue development during the 3 stocker/backgrounding phase of production. During the stocker/backgrounding phase of production, significant muscle growth occurs and the primary structures of marbling deposits are developed. Although increasing intramuscular fat deposition relative to other fat depots would be beneficial, little is known about the association between muscle growth, metabolism and marbling development. Fat Cell Differentiation and Development Understanding the molecular mechanisms responsible for fat cell development may provide insight into how fat deposition can be influenced during the stocker phase of production, whether from nutrition, management, or genetics. Growth of intramuscular fat appears to occur due to adipocyte (fat cell) acquisition by differentiation of preadipocytes (adipogenesis; Minoshima et al., 2001). Adipocytes represent between 1/3 and 2/3 of the cells in adipose tissue. The remaining 1/3 to 2/3 consists of blood cells, endothelial cells, pericytes, preadipocytes, and fibroblasts (Ailhaud et a1., 1992). Late in embryonic life the first preadipocytes appear. However, expansion of white adipocytes does not occur until shortly after birth (Slavin, 1979; Cook and Kozak, 1982). The expansion of white adipocytes that occurs shortly after birth includes both an increase in the size (hypertrophy) and number (hyperplasia) of adipose cells. Interestingly, even in adult animals, the potential to increase the number of adipose cells exists (Gregoire et al., 1998). Lineage of adipose is derived from multipotent stem cells of mesodermal origin. These multipotent stem cells can differentiate into muscle, cartilage, as well as fat (Cornelius et al., 1994). Adipogenic cells, as well as osteogenic and chondrogenic cells, may arise from sclerotomal cells (Gregoire et al., 1998). Adipoblasts formed from multipotent stem cells undergo commitment to form preadipocytes. Preadipocytes are characterized as possessing early genetic markers of adipocytes, but they have not at this point accumulated triacylglycerol 4 (lipid) stores (Ailhaud et al., 1992). The process of adipose differentiation corresponds to phenotypic changes beginning with the adipoblast, progressing to the preadipocyte, becoming an immature fat cell, and finally, terminally differentiating into the mature fat cell (Ailhaud et al., 1992). In order to induce the differentiation of preadipocytes to mature adipocytes, external inducers are required. Inducers considered to be required for adipocyte differentiation to occur include insulin-like growth factor-1 (IGF-1), glucocorticoid, cyclic adenosine monophosphate (cAMP), and long chain fatty acids. Considerable numbers of IGF-1 receptors are found on preadipocytes. In addition, insulin and growth hormone, driven by nutrition and management (e.g., anabolic implants), are able to induce adipocyte differentiation via the IGF-l receptor. Glucocorticoid is an adipogenic agent required for preadipocytes to undergo clonal expansion and terminal differentiation (MacDougald and Lane, 1995). In addition, increased levels of cAMP are required for preadipocyte cells to undergo differentiation into mature adipocytes (Vassaux et al., 1992). It has also been shown that long chain fatty acids can induce differentiation of preadipocytes. For example, palmitic acid (C16:0) can induce growth arrested cells to undergo post-confluence mitosis (clonal expansion), to accumulate triacylglycerol, and to express several adipocyte markers (Amri et al., 1995). Intramuscular fat cells develop within the perimysium of muscle bundles in close association with the skeletal muscle satellite cells responsible for postnatal muscle growth. Therefore, skeletal muscle growth and metabolism might influence marbling development. Intramuscular fat content is positively correlated (r = 0.46 to 0.49) with fiber area of oxidative muscle fiber types (Melton et al., 1974; May et al., 1977) and number of capillaries per muscle fiber (r = 0.63; Melton et al., 1975). In addition, Kim et al. (2009) reported that gene expression 5 of NADH dehydrogenase and cytochrome c oxidase, both involved in oxidative phosphorylation within mitochondria, were greater in muscle from the loin, which had greater intramuscular fat content, compared with muscle from the top round. Gene expression of NADH dehydrogenase and cytochrome c oxidase explained 84 and 97% of the variation in intramuscular fat content of loin muscle, respectively, in Korean cattle (Kim et al., 2009). These data suggest that muscle metabolism influences marbling development. Possible mechanisms by which muscle influences marbling development could include increased angiogenesis (growth of blood vessels) within muscle providing greater blood flow to sustain new adipocytes and changes in intercellular signaling mechanisms such as growth factors that stimulate adipocyte differentiation. Angiogenesis is an important aspect of adipose tissue development, and adipocyte differentiation may be regulated by factors that stimulate angiogenesis (Hausman and Richardson, 2004). Under the microscope, it can be seen that intramuscular adipocytes develop in close proximity to a capillary network (Harper and Pethick, 2004), and an important player in angiogenesis, the plasminogen activator-plasmin system, is involved in the migration of endothelial cells and preadipocytes (Hausman and Richardson, 2004). Thus, enhancing vascular development of muscle tissue could stimulate recruitment of preadipocytes to new locations within the muscle forming new intramuscular fat deposits. As compared to other adipose tissues, intramuscular adipose tissue is more likely to be influenced by the environment within the muscle tissue. Intramuscular adipose tissue has smaller mature adipocytes (Smith and Crouse, 1984; Cianzio et al., 1985; May et al., 1994) and preadipocytes from intramuscular adipose tissue have reduced capacity to differentiate (Grant et al., 2008; Ortiz-Colon et al., 2009) compared to other adipose tissue depots. This may be due to effects of the intramuscular environment or differences in the genetically determined growth 6 potential. Postnatal growth of bovine adipose tissue occurs by hypertrophy (cell size) through fat synthesis, and hyperplasia (cell number) due to differentiation of preadipocytes (Gerrard and Grant, 2003). As mentioned, several studies (Smith and Crouse, 1984; Cianzio et al., 1985; Schoonmaker et al., 2004) have reported that intramuscular adipose tissue has a greater number of smaller adipocytes compared to subcutaneous and perirenal adipose tissue. In addition, perirenal adipose tissue has greater mean adipocyte diameter than subcutaneous adipose tissue (Prior, 1983; Cianzio et al., 1985). These differences in hypertrophy may be related to rate and substrate utilized for triacylglycerol synthesis. Smith and Crouse (1984) reported that intramuscular adipose tissue had greater rates of fatty acid synthesis from glucose compared to subcutaneous adipose tissue, whereas, subcutaneous adipose tissue had greater rates when acetate or lactate were provided in vitro. However, rates of glycerol synthesis were greater for subcutaneous adipose tissue irrespective of carbon source suggesting that subcutaneous adipose tissue has greater rates of triacylglycerol synthesis and lipid storage than intramuscular adipose tissue. Results of Rhoades et al. (2007) also indicated that rates of glycerol synthesis were greater in subcutaneous than intramuscular adipose tissue. In addition, glucose incorporation into lipids in intramuscular adipose tissue responded to insulin, whereas insulin had no effect in subcutaneous adipose tissue (Rhoades et al., 2007). The response of intramuscular adipose tissue was also modulated by diet, such that a corn-based high-glucose diet resulted in an insulin response, whereas, a hay-based low-glucose diet did not. Differences in insulin response related to diet may also exist between subcutaneous and internal fat depots. Developmental differences between intramuscular and other fat depots have lead to an interest in nutrition and management strategies to enhance marbling deposition while decreasing fat deposition in other depots. Because intramuscular fat deposition is a lifetime event and 7 potentially associated with muscle growth and development, stocker programs that vary in nutrient quantity and quality and cattle growth rates likely have a significant impact on marbling deposition. FACTORS THAT IMPACT MARBLING IN STOCKER CATTLE Several factors have been shown to alter carcass quality in stocker cattle including animal health, nutrition, management and genetics. Animal Health Bovine Respiratory Disease. Verification of health or preconditioning programs for feeder calves before they begin the finishing phase of production is becoming more common in the beef cattle industry. With the demand for higher quality products, the increase in valuebased marketing, and the increase in vertical coordination systems, both cow-calf producers and feedlot operators have become more in-tune to health management practices that have the potential to increase overall profitability. The health status of calves upon arrival to the feedyard has been shown to impact the efficiency of cattle in the feedyard, and also to affect the quality attributes of the cattle at slaughter. McNeill (1994, 1995, 1996, 1997, 1998) and Montgomery et al. (1984) documented that morbid cattle during the finishing phase of production had carcasses with a lower degree of marbling and subsequently lower USDA Quality Grades. Therefore, although the medical costs attributable to the treatment of bovine respiratory disease (BRD) are substantial (Martin et al., 1982; Perino, 1992), the economic impact of BRD on animal performance, carcass merit, and meat quality are likely even more devastating. Morbidity rates account for approximately eight percent of all production costs without consideration of losses due to decreased performance (Griffen et al., 1995). McNeill et al. 8 (1996) reported that “healthy” steers had greater daily gains and 12% more USDA Choice carcasses than cattle identified as “sick” at some point during the finishing period. Gardner et al. (1999) showed that steers with lung lesions plus active lymph nodes had $73.78 less net return, of which 21% was due to medicine costs and 79% due to lower carcass weight (8.4% less) and lower quality grade (24.7% more USDA Standards). This negative impact on carcass traits 200 d after receiving the cattle illustrates the importance of preventing BRD. Roeber et al. (2001) documented that although cattle treated more than once in the feedyard had lower marbling score and hot carcass weight (HCW) than those not treated, within a quality grade class, no differences for tenderness or palatability traits existed in cattle stratified by number of hospital visits or preconditioning treatment program. Garcia et al. (2010) evaluated the effects of BRD on carcass composition and meat quality traits in two separate herds at the U.S. Meat Animal Research Center, Clay Center, NE. Cattle showing signs of BRD experienced a 11 lb decrease in HCW, and a 10 to 15% decrease in 12th-rib fat thickness resulting in a 5 to 7% decrease in USDA Yield Grade. These authors also evaluated the effects of overall pathogenic disease on carcass traits. If an animal experienced any disease event, 12th-rib fat thickness was decreased 12 to 14%, and USDA Yield Grade was decreased by 0.2 units. Similarly, Schneider et al. (2009) reported a 2.5% decrease in HCW, a slight decrease in LM area, a 7.0% decrease in 12th-rib fat thickness, and a 2.5% decrease in marbling score in treated vs. non-treated cattle. Gardner et al. (1999) examined the effects of the number of times treated on carcass characteristics. Hot carcass weight was decreased 4.1%, 12th-rib fat thickness was decreased 26.5%, and KPH and USDA Yield Grade were also decreased in treated vs. non-treated calves. Similarly, HCW was decreased by 4.7%, 12th-rib fat thickness by 43.4%, and KPH and USDA Yield Grade by 21.1 and 18.2%, respectively, when 9 cattle were treated more than once compared with cattle only treated once (Gardner et al., 1999). When morbidity was defined by the presence of lung lesions at slaughter, Gardner et al. (1999) observed morbid steers had HCW that were 94% of HCW of steers with no evidence of prior disease. Additionally, when active bronchial lymph nodes were present, indicating an ongoing disease process, HCW were less than HCW of animals with no lesions. In general, evidence of prior morbidity was associated with decreased dressing percentage, internal fat, external fat, smaller LM area, and lower marbling scores. Similar results were noticed when carcass characteristics were examined based on BRD treatment records. Cattle that received more than one treatment for BRD had an additional 5% decrease in HCW compared with those that were treated only one time (Gardner et al., 1999). Roeber et al. (2001) reported a 3% decline in HCW in steers treated more than one time for BRD compared to those treated one time. However, in their report, no differences in HCW were observed between cattle never treated and those treated one time, and no differences in HCW across number of BRD treatments were reported by Waggoner et al. (2007). Although fewer differences in carcass weights and no differences in LM area were reported by Roeber et al. (2001), measures of carcass fatness were affected by BRD incidence. While internal fat was not different among treatment groups, 12th-rib fat thickness, calculated yield grade, and marbling score were all greatest for cattle never treated, intermediate for those treated once, and lowest after multiple treatments. As a result of differences in marbling, the distribution of USDA Quality Grades has been shown to be affected by BRD. Gardner et al. (1999) observed that treated steers, or steers with lung lesions, had a greater percentage of USDA Standard carcasses at the expense of Choice and Select compared with healthy animals. Similarly, McNeill et al. (1996), in an evaluation of over 7,000 cattle, reported 39% of cattle never treated 10 for BRD graded USDA Choice or better, but only 27% that were treated graded USDA Choice or better. Holland et al. (2010) noted a trend for a linear decrease in marbling score with increasing number of treatments for BRD when cattle were fed to a targeted 12th-rib fat thickness of 1.27 cm. There were no differences in proportions of USDA Yield Grades or Quality Grades; however, meat color score and overall appearance of strip loin steaks decreased as the number of times treated for BRD increased. There was no effect of number of BRD treatments on Warner Bratzler Shear Force (WBSF), but there was a trend for an increased amount of connective tissue determined by trained sensory panelists as the number of treatments increased. It is apparent that BRD has negative effects on carcass characteristics. Data show that fat deposition by cattle is the most directly affected; however, decreased growth and anorexia also occurs as evidenced by decreased HCW. It may be possible to achieve similar HCW, although animals would need to be sorted by number of times treated for BRD and fed for additional days (Holland et al., 2010). Other than in chronically infected cattle, this would indicate that BRD constraints on growth are relieved with time. However, the trend for decreased marbling score with increased number of treatments observed by Holland et al. (2010) suggests that quality of carcasses may not fully recover even with increased days on feed. Deworming. Internal parasites in cattle have been shown to decrease performance and impair immune function (Wiggin and Gibbs, 1990; Smith et al., 2000; Reinhardt et al., 2006). In feedlot cattle, infection of cattle by Ostertagia ostertagi decreased feed intake, average daily gain (ADG), and efficiency of protein digestion (Fox et al., 1989), and infected cattle treated with fenbendazole ate more feed than nontreated controls (Fox et al., 1989; Smith et al., 2000). Reinhardt et al. (2006) reported that the percentage of USDA Prime or Choice carcasses tended 11 to be increased and USDA Standard carcasses decreased when feedlot heifers were treated with fenbendazole and ivermectin pour-on compared to heifers treated with ivermectin pour-on alone. Marbling score also tended to be greater for heifers treated with the combination of fenbendazole and ivermectin pour-on. The increased degree of body fat was consistent with the noted increase in dry matter intake (DMI). Feedlot deworming of steers not dewormed on pasture increased carcass weight by 49 lb, while feedlot deworming of steers that were dewormed on pasture increased carcass weight by 21 lb (Smith et al., 2000). The percentage of Choice carcasses was lower and the percentage of Select carcasses was greater for steers not dewormed on pasture or in the feedlot compared to treatments groups that were dewormed. Deworming during grazing resulted in a net benefit of $33.75 per steer had steers been sold at the end of the grazing phase. For the complete grazing-finishing system, feedlot deworming of previously non-dewormed steers with fenbendazole produced a net benefit of $30.61 per steer on a carcass-adjusted basis, while feedlot deworming of dewormed steers produced a net benefit of $11.07. Therefore, there are clear performance and economic benefits to deworming grazing steers on pasture, and to deworming yearling steers entering the feedlot from summer pasture. In general, management practices that promote improved animal health will have a positive impact on marbling score and carcass quality. Nutrition As previously mentioned, Smith and Crouse (1984) reported that intramuscular adipocytes preferentially utilize glucose as the primary substrate for fatty acid synthesis; whereas subcutaneous fat utilizes acetate. This led to studies conducted at the University of Illinois where researchers showed that suckling calves provided a corn-based creep feed had increased marbling scores at the end of finishing compared to calves provided a soyhull-based creep feed, 12 suggesting that increased glucose supply from corn supplementation improved marbling deposition (Faulkner et al., 1994). Similarly, Myers et al. (1999b) and Shike et al. (2007) reported that early weaned calves placed directly on a finishing diet had greater marbling scores compared with calves that were normal weaned with and without creep feed. These data suggest that an increase in starch intake early in an animal’s life may begin to initiate marbling development and improve final carcass quality. Numerous investigations have also sought to determine the effects of nutrition and management throughout the production chain on deposition of marbling and subsequent carcass quality (Berger and Faulkner, 2003). Increases in marbling have been shown by starting cattle on high-concentrate rations at earlier ages (Myers et al., 1999a; 1999b; Wertz et al., 2001, 2002); however, these cases dealt with calves that were early weaned and started on feed at very early ages (70 to 100 days of age). Wertz et al. (2002) evaluated the effect of weaning calves and growing them on forage compared to early-weaning calves and immediately placing them on a high-concentrate diet. Early-weaned heifer calves were approximately 20% more efficient at any ultrasound marbling score compared with older heifers that had grazed and were placed on feed later in life. In addition, at any given 12th-rib fat thickness, early-weaned calves had higher ultrasound marbling scores. In a similar study, Angus × Simmental heifer calves that were fed high-energy diets at 208 days of age or earlier deposited more marbling relative to 12th-rib fat than heifers of the same genetics that were finished as long yearlings (Wertz et al., 2001). In normal-weaned calves (approximately 200 days of age), Faulkner et al. (1994) showed that creep feed energy source could affect carcass quality even when weight gain was similar during the creep phase. In their experiment, calves which had been fed a corn-based creep had higher marbling scores than calves which had been creep fed soy hulls. In contrast, no difference 13 in marbling score was observed in normal-weaned calves creep-fed high corn, high fiber, or no creep (Berger and Faulkner, 2003). Reasons for discrepancies are unclear, although Bruns (2006) suggested that if calves can sustain their normal growth curve without additional supplementation then creep feeding won’t increase quality grade. However, if the calves are growing below their growth potential, due to inadequate nutrition, then creep feeding or early weaning should help their ability to grade. Even though early weaned calves generally have lower average daily gains in the finishing phase than yearlings, they are consuming excess calories early in life above their requirement for normal growth. Therefore, marbling (intramuscular fat) deposition might be greater compared with normal-weaned calves (Bruns, 2006). Level of dietary starch during backgrounding. We conducted a meta-analysis to evaluate the level of dietary starch in growing diets during the stocker/backgrounding phase. A dataset with a total of 14 studies was compiled comparing growing diets fed to beef cattle in the drylot that differed in grain content of the diet (i.e., starch content) for 56 to 145 d prior to finishing. To be included in the dataset, feed intake of the different diets had to be adjusted to provide similar NEg intake and achieve similar rates of gain during the growing phase. Treatments were categorized as either low (LS), medium (MS), or high (HS) starch diets based on grain and NEg content of the diet. Corn was the predominate cereal grain used except in the study by Sainz et al. (1995), which used rolled wheat in the HS diet. Grain content of the diet was calculated from the reported ingredient composition of the experimental diets, where corn or sorghum silage was assumed to contain 50% grain. Seven studies had a comparison of HS versus MS, and 9 studies had a comparison of HS versus LS. Only 2 studies had a comparison of MS versus LS; therefore this comparison was not analyzed. The dataset was divided into two sub-datasets of either HS 14 versus MS or HS versus LS. Grain content, NEg concentration, and growing ADG for HS versus MS, and HS versus LS sub-datasets are presented in Table 1. Each sub-dataset was analyzed using a linear mixed model (Proc Mixed of SAS, SAS Instit., Cary, NC) that included diet as a fixed effect and intercept as a random effect with the unstructured option used for the var-(co)var matrix and trial as the subject. Least squares means were computed using the inverse of the squared standard error for the dependent variable as a weighting factor (St-Pierre, 2001). Least squares means were compared using Tukey’s W procedure and were considered different at P < 0.10. Results of the HS versus MS comparison are presented in Table 2. Finishing performance was similar between diet types with no differences for ADG, DMI, or gain:feed. In addition, there were no differences in LM area, rib fat thickness, KPH, yield grade, or marbling score between steers previously fed HS or MS diets during the growing phase. Of the individual studies used in the meta-analysis, the results consistently show little to no difference in marbling score with regard to grain content of the growing diet. In three trials, Loerch (1990) fed Angus crossbred steers a corn silage-based growing diet ad libitum or high moisture corn-based diet at restricted intake for 85 d prior to finishing on a common diet. In all three trials, growing diet had no impact on final marbling score. Similarly, Sip and Pritchard (1991), Coleman et al. (1995) and McCurdy et al. (2010) found that corn-grain based growing diets did not improve marbling score compared with corn/sorghum silage-based growing diets. Gunter et al. (1996) and Vasconcelos et al. (2009) reported that marbling scores were similar between steers fed 90% concentrate or 50 to 60% concentrate growing diets prior to finishing on a common diet. When comparing finishing performance of steers fed HS or LS diets during the growing phase, there were no differences in final BW, DMI, or gain:feed; however, steers previously fed 15 HS diets had greater (P < 0.10) ADG (1.76 vs. 1.67 kg/d, respectively) during the finishing phase (Table 3). Steers fed HS or LS diets during the growing phase had similar LM area, rib fat thickness, KPH, yield grade, and marbling score. Wagner (1988) fed growing steers a corn silage/alfalfa-grass hay diet ad libitum or high moisture corn diet limit-fed to provide similar ME intake as the corn silage/alfalfa-grass hay diet. Following the finishing phase, marbling score was similar between growing diets. Similarly, Sainz et al. (1995) and Hill et al. (1996) observed that final marbling score was similar between steers fed an alfalfa hay/cottonseed hull-based growing diet or limit fed a grain-based diet. Vasconcelos et al. (2009) reported that steers limit fed a steam-flaked corn diet had no impact on final marbling score compared with steers fed a wheat midd/cottonseed hull growing diet ad libitum. McCurdy et al. (2010) compared carcass characteristics of steers limit fed a grain-based diet with steers that grazed winter wheat pasture managed such that rate of gain during the growing phase was similar (1.18 vs. 1.15 kg/d, respectively). After finishing on a common diet, marbling score was similar between steers that grazed wheat pasture and those that were fed the grain-based growing diet. Collectively, metaanalysis of these studies indicate that dietary starch content of growing diets has little impact on final marbling score. Level of starch supplementation during grazing. Supplementation of starch to grazing cattle during the stocker phase has received less attention due to limited ability to dramatically change the starch content of the overall diet compared with growing diets fed in drylot. However, a few studies have evaluated starch-based supplements to grazing stocker cattle on final carcass quality. Horn et al. (1995) reported that high-starch corn-based supplements fed to steers grazing winter wheat pasture did not improve marbling score compared with steers fed a high-fiber soybean hull/wheat midd-based supplements. Similarly, Bumpus (2006) and Sharman 16 et al. (2012) observed that high-starch supplements did not improve marbling score of steers grazing ryegrass and winter wheat pasture, respectively, compared with high-fiber supplements. Sharman et al. (2009) fed either corn, soybean hulls, or distillers grains at 1% of BW to steers grazing dormant winter native range. Similar to previous studies, these authors observed no difference in final marbling score between the different supplement types, even though glucose supply for steers grazing dormant native range would have been very low. Interestingly, Lake et al. (1974) and Lomas et al. (2009) reported that supplementing steers grazing cool-season pasture with 1.82 or 1.64 kg/d of grain increased marbling score compared with no supplementation. However, in a second trial, Lake et al. (1974) observed no improvement in quality grade when steers were supplemented up to 2.72 kg/d of corn. The explanation for these differences could be that the starch-based supplement was not matched with a fiber-based supplement such that the supplement provided additional energy for fat deposition rather than a starch effect. Similarly, Bumpus (2006) observed that starch- and fiberbased supplements improved marbling score compared with non-supplemented steers, but rate of gain during the stocker phase was not increased for supplemented steers compared with nonsupplemented steers. In contrast, Horn et al. (1995), Sharman et al. (2009), and Sharman et al. (2012) observed that starch- or fiber-based supplements did not improve marbling score compared with non-supplemented steers. Therefore, simply the additional energy does not appear to explain the improved marbling scores. Another explanation could be the BW of cattle during the stocker phase relative to mature weight. Carter et al. (2002) evaluated the relationship between ultrasound intramuscular fat percentage and BW when steers were grown on pasture and placed on feed at three different weights (363, 408, or 454 kg initial finishing BW). These authors observed that regardless of 17 placement weight there was breakpoint BW of 378 kg at which intramuscular fat began to increase. The breakpoint BW of 378 kg was calculated to be 64% of mature weight of the steers or 66% of final feedlot BW. When applying this concept to previous studies, feedlot placement BW of steers in Trial 1 of Lake et al. (1974), Bumpus (2006), and Lomas et al. (2009) were 73, 70, and 70% of final feedlot BW indicating that energy supplementation had increased energy intake after the point at which intramuscular fat deposition increased. In contrast, feedlot placement BW was only 66, 62, and 54% of final feedlot BW in the studies of Lake et al. (1974; Trial 2), Horn et al. (1995), and Sharman et al. (2009) where final marbling score was not influenced by energy supplementation during the stocker phase. In conclusion, the starch content of growing diets or energy supplements fed to grazing cattle does not appear to influence final marbling scores. In addition, energy supplementation of lightweight stocker cattle may not improve final marbling score. However, energy supplementation of heavyweight yearling stocker cattle could be beneficial for enhancing final marbling score of cattle. Further research is necessary to validate the concept that providing an energy supplement to heavyweight yearling stocker cattle could improve marbling scores. Rate of gain during the stocker phase and placement weight. A second dataset consisting of 29 trials was compiled to evaluate the relationships of rate of gain during the stocker/backgrounding phase and initial BW at start of finishing with carcass characteristics. To be included in the dataset, normal-weaned steers/heifers must have entered the stocker phase shortly after weaning, and stocker treatments must have differed in rate of gain by > 0.10 kg/d during the stocker phase. In this dataset, steers/heifers either grazed different forage types, were fed with different levels of supplement while grazing pasture, or were fed high-roughage growing diets in drylot to achieve different rates of gain during the stocker/backgrounding phase 18 prior to being fed a common finishing diet. In some trials, treatments were fed for similar days on feed during the finishing phase, whereas in other trials treatments were fed to a common fat endpoint with different days on feed. Mixed model regression analysis following random coefficient methodology (St-Pierre, 2001) was used to evaluate the relationships of rate of gain during the stocker/backgrounding phase and initial finishing BW with carcass characteristics and rib fat-adjusted carcass characteristics. A general linear model (Proc GLM of SAS) that included the covariate (i.e., stocker ADG or initial BW), trial, and trial x covariate interaction terms was used to test the hypothesis that the slope between the covariate and carcass characteristics was similar among trials. Regression analyses were conducted using a mixed model (Proc Mixed of SAS) that included the covariate(s) as fixed effects, and intercept and covariate(s), when slopes differed among trials, as random effects with the unstructured option used for the var-(co)var matrix and trial as the subject. Regression coefficients were computed using the inverse of the squared standard error for the dependent variable as a weighting factor (St-Pierre, 2001), and were considered different from zero at P < 0.10. Quadratic regression coefficients were tested and were not significant. Summary statistics for stocker performance and carcass characteristics of the 29 trials are presented in Table 4. Stocker phase ADG and initial finishing BW were positively related with HCW, LM area, and KPH, but not rib fat thickness or yield grade (Table 5). Hot carcass weight was positively related to LM area, rib fat thickness, and yield grade, but not KPH. These results indicate that as cattle grew faster during the stocker phase and were heavier entering the feedyard they produced heavier carcasses with larger longissimus muscle area, but had no influence on carcass fat composition as evidenced by the lack of relationship with fat thickness and yield grade. 19 Marbling score was not related to stocker phase ADG (Figure 1) or initial finishing BW (Figure 2), but was positively related to HCW (Figure 3) suggesting that growth during the stocker phase has little influence on marbling score and it is the weight of the animal at slaughter that has the largest influence. Additionally, marbling score was positively related to rib fat thickness [y = 291.82 ± 29.2048 + 92.5465 ± 22.2935 * rib fat thickness (cm)]. Given the relationship between marbling score and rib fat thickness and to evaluate relationships with rib fat-adjusted carcass characteristics, rib fat thickness was included in the regression of stocker phase ADG, initial finishing BW, and HCW with marbling score. When adjusted for rib fat thickness, marbling score was positively related with stocker phase ADG, initial finishing BW, and HCW suggesting that all three factors may play a role in influencing marbling score when cattle are fed to a similar rib fat thickness. To determine the relative importance of stocker phase ADG, initial finishing BW, and HCW with rib fat-adjusted marbling score, these traits were included in the regression model to predict rib fat-adjusted marbling score. When stocker ADG, initial finishing BW, and HCW were included in the model along with rib fat thickness, HCW continued to be positively related to marbling score, but stocker phase ADG and initial finishing BW were no longer related to marbling score (Figures 1, 2, and 3, respectively). Thus, it appears that increasing HCW will increase rib-fat adjusted marbling score regardless of previous management. Our regression analysis indicates that greater stocker phase ADG (Figure 4) or initial finishing BW (Figure 5) can increase HCW, which can be accomplished through the use of higher quality forage, energy supplements, or longer grazing periods. However, to increase rib fat-adjusted HCW, regression analysis indicates that lower rates of gain (Figure 4) and longer grazing periods to achieve greater initial finishing BW (Figure 5) are needed. 20 These results may explain the lack of differences in marbling score in cattle from different stocker and backgrounding studies (Coleman et al., 1995; Sainz et al., 1995; Hersom et al., 2004; McCurdy et al., 2010), because even though cattle in these studies underwent widely different growing programs the end result was similar HCW when fed to similar rib fat thickness. Our results are also contrary to the idea that cattle should be grown at moderate to high rates of gain and finished at young ages to achieve high quality grades. In agreement with our results, Klopfenstein et al. (2000) reported that long-yearling production systems resulted in greater initial finishing BW and HCW, and increased the percentage of cattle grading choice when adjusted to similar rib fat thickness compared with calf-fed production systems. Similarly, Gill et al. (1993) and Sharman et al. (2012) observed that steers wintered on dormant native range followed by season-long grazing of native pasture prior to finishing increased initial finishing BW and tended to increase marbling scores but lower rib fat thickness compared with steers grazing winter wheat pasture prior to finishing. In conclusion, marbling score and rib fat thickness-adjusted marbling score can be manipulated using nutrition and management during the stocker phase. Marbling scores can be improved by ‘making cattle bigger’ through increasing rate of gain during the stocker phase, initial finishing BW, and HCW. However, this will also increase rib fat thickness and yield grade resulting in little improvement in marbling to rib fat ratio. In contrast, rib fat-adjusted marbling score (i.e., improved ratio of marbling to rib fat) can be improved by using low to moderate rates of gain for longer grazing periods during the stocker phase to increase initial finishing BW and rib fat-adjusted HCW. Management 21 Growth Promoting Implants. Anabolic implants increase weight gains of beef cattle by 10 to 20% and are one of the most profitable management tools available to beef producers. However, use of anabolic implants has been shown to have a negative impact on quality grade. Duckett et al. (1997) reported that estrogenic implants administered to steers during finishing decreased marbling score from 26 to 44 points; however, androgen alone had less effect, decreasing marbling score 22 points and being statistically similar to controls. Morgan (1997) reported that implants containing estrogen alone reduced marbling score 12 to 47 points, androgen alone reduced marbling score 0 to 9 points, and combination implants reduced marbling score 12 to 29 points in feedlot steers and heifers. In a meta-analysis of published studies, McPhee et al. (2006) reported that estrogenic or combination implants reduced intramuscular fat percent by 0.32 percentage units. Reinhardt (2010) reported that a metaanalysis of feedlot implant trials showed a reduction in marbling score of 49 points (548 vs. 499) when using a very aggressive implant strategy of multiple strong combination implants. Reinhardt (2010) also reported that as the implant program became more aggressive the decrease in marbling score compared to non-implanted cattle became greater. The reason for decreased marbling scores of implanted cattle could be attributed to a shift in energy use for muscle growth rather than fat deposition or changes in the intramuscular environment directing progenitor cells toward myogenesis rather than adipogenesis. Anabolic implants alter body composition by decreasing fat composition and increasing lean. The decrease in marbling score could be due to reduction in total fat deposition, but many studies (Simms et al., 1988; Mader et al., 1994; Platter et al., 2003; Scheffler et al., 2003) have reported reduced marbling scores even though cattle are fed to similar rib fat endpoints. Thus, the impact of implants on marbling score appears to be due to a specific effect on intramuscular fat. In a 22 review, Chung and Johnson (2007) described a plausible mechanism whereby anabolic implants could be stimulating progenitor cells within skeletal muscle to follow a myogenic lineage rather than an adipogenic lineage. Thus, intramuscular fat development is influenced to a greater degree than other fat depots. There is concern that implanting cattle earlier in the production cycle (suckling or stocker phase) has further negative impacts on quality grades. Platter et al. (2003) reported that backgrounding implant tended (P =0.09) to decrease marbling score by 25 points. These authors also reported that steers receiving 4 or 5 implants during their lifetime had significantly reduced marbling scores (38 points) compared to steers only implanted twice in the feedlot. McMurphy et al. (unpublished data) observed that fall-born steers implanted with Component TE-G after weaning tended (P = 0.09) to have marbling scores 32 points lower than those not implanted during the stocker phase. In contrast, Sharman et al. (2011) observed that fall-weaned steers implanted with Component TE-G during the stocker phase had similar marbling scores compared to steers not implanted during the stocker phase. The different responses observed in these studies may be due to forage quality which would change energy available for fat deposition. In the study by McMurphy and others, steers grazed summer native range, whereas steers grazed winter wheat pasture in the study by Sharman et al. (2011). However, Paisley et al. (1999) reported that steers implanted with Synovex-C, Synovex-S, or Revalor-G while grazing lowquality dormant winter native range had similar marbling scores compared with non-implanted steers suggesting that forage quality does not influence the effect of stocker phase implants on quality grades. Several studies (McCann et al., 1991; Brandt et al., 1995; Fankhauser et al., 1997) have reported no effect of implants during the stocker phase on carcass quality grades. In addition, Mader et al. (1985) and Simms et al. (1988) reported that steers implanted during the 23 growing phase while fed a corn silage-based diet in drylot had similar marbling scores compared to steers implanted only during the finishing phase. Mader et al. (1994) observed that implanting steers with Synovex-C during the suckling phase and Synovex-S during the stocker and finishing phase did not negatively impact marbling score compared with steers only implanted during the stocker and finishing phases. Few studies have evaluated the effect of type of implant in stocker cattle on carcass characteristics. Kuhl et al. (1997) evaluated the use of Ralgro, Revalor-G and Synovex-S for grazing steers and reported no differences in carcass quality traits or yield grade among implants. Frankhauser et al. (1997) evaluated Ralgro and Synovex-S in steers grazing native pastures under intensive early stocking management prior to finishing. Type of grazing implant did not influence marbling score or yield grade. Paisley et al. (1999) administered Synovex-C, SynovexS, or Revalor-G to steers grazing dormant winter native range prior to being implanted with Ralgro while grazing summer pasture until mid-July. Following the subsequent finishing phase, steers implanted with Synovex-C tended (P = 0.06) to have greater marbling scores than steers implanted with Synovex-S or Revalor-G. Possibly the lower potency of Synovex-C did not negatively affect intramuscular fat development; however, the non-implanted steers had marbling scores intermediate to Synovex-C and Synovex-S/Revalor-G steers making it difficult to discern the cause of the greater marbling score of Synovex-C steers. Interestingly, Scheffler et al. (2003) and Simms et al. (1988) observed that the terminal implant administered late in the finishing phase had the greatest negative impact on marbling score. Scheffler et al. (2003) implanted long-fed Holstein steers 1 time on d 224, 2 times on d 112 and 224, 3 times on d 0, 112, and 224, or non-implanted control during a 291-d finishing period. These authors reported that the single implant on d 224 decreased marbling score by 52 24 points compared to non-implanted controls, whereas implanting steers 2 or 3 times throughout the finishing period only reduced marbling score by 31 and 34 points, respectively. Simms et al. (1988) evaluated different combinations of four implant timings during the suckling, growing, and finishing (early and late) phases compared with non-implanted steers. These authors reported that any combination of implanting steers during the production cycle that included the latefinishing implant at 75 d prior to harvest decreased quality grade compared to non-implanted controls. However, a combination of implants that eliminated the late-finishing implant did not significantly decrease quality grade compared to non-implanted steers even though this combination of implants included suckling, growing, and early-finishing implants. Therefore, it appears that administration of implants closer to the end of the finishing phase have a greater negative impact on quality grades, whereas administration of implants early in the production cycle have little influence on quality grades if cattle are to be implanted during the finishing phase. Genetics Bertrand et al. (2001) reported heritability estimates for several carcass traits adjusted to an age constant or time-on-feed constant basis. The average heritability estimates for all traits were in the moderate-to-high range, although there was a higher frequency of low (≤ 0.12) estimates observed for Warner-Bratzler shear force and tenderness score. Large genetic differences across breeds and small genetic differences within the breeds have been reported for shear force and taste panel tenderness scores from reciprocal backcross and single-cross steers of several breeds (Van Vleck et al., 1992). Van Vleck et al. (1992) also reported that large acrossbreed and within-breed genetic variation existed for marbling score. Adequate within-breed genetic variation for shear force might warrant its inclusion in within-breed genetic evaluation 25 programs for breeds of cattle (Bertrand et al., 2001). In the review by Bertrand et al. (2001), heritability estimates for carcass weight, ribeye area, shear force, marbling score, and calpastatin activity averaged across studies evaluating carcasses at a fat constant endpoint were also shown to be moderate to high. The authors concluded that on average, heritability estimates shown in the literature indicate that carcass traits will respond well to selection, and that it may be possible to select for increased growth within a breed without negatively impacting marbling score or percentage retail product. Quantitative trait loci (QTLs) are stretches of DNA that are closely linked to the genes that underlie any given trait, such as characteristics of carcass quality and yield. Quantitative trait loci can be molecularly identified to help “map” regions of the bovine genome that contain genes involved in specifying a given quantitative trait. Bertrand et al. (2001) suggested that the QTL that are identified through searches of the bovine gene map are likely to be most beneficial for those traits that are difficult and/or expensive to measure, such as disease resistance and immunocompetence, carcass quality and palatability attributes, fertility and reproductive efficiency, maintenance requirements (i.e., energetic efficiency), carcass quantity and yield, milk production and maternal ability, and growth performance (ranked from greatest benefit to least benefit). Because hard-to-measure traits such as immunocompetence have a significant effect on carcass quality, DNA-based tools for carcass trait selection will become increasingly useful. CONCLUSIONS No one factor solely contributes to carcass quality, but numerous factors interact to have an effect. Calf health, nutrition, and management (e.g., implant strategy) prior to and during feedlot finishing can have an impact on carcass quality. It is important to maintain the genetic potential of calves to express growth performance and carcass quality through health 26 management programs and implant strategies during the stocker phase. Adequate caloric intake at critical times to support both muscle and marbling development appears to improve the percentage of cattle that will grade USDA Choice. Marbling scores can be improved by ‘making cattle bigger’ through increasing rate of gain during the stocker phase, initial finishing BW, and HCW. However, this will also increase rib fat thickness and yield grade resulting in little improvement in marbling to rib fat ratio. In contrast, rib fat-adjusted marbling score (i.e., improved ratio of marbling to rib fat) can be improved by using low to moderate rates of gain for longer grazing periods during the stocker phase to increase initial finishing BW and rib fatadjusted HCW. 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Adverse immune reactions and the pathogenesis of Ostertagia ostertagi infections in calves. Am. J. Vet. Res. 51:825–832. 41 Table 1. Mean (Range) for grain content, NEg concentration, and growing phase ADG for steers fed high versus medium starch, or high versus low starch diets during the growing phase Dataset 1 Dataset 2 Variable High Starch Medium Starch High Starch Low Starch Grain, % 79.4 (70.8 – 89.2) 45.2 (35.0 – 52.0) 72.8 (65.0 – 79.2) 16.4 (0.0 – 35.9) NEg, Mcal/kg DM 1.42 (1.21 – 1.54) 1.02 (0.62 – 1.22) 1.42 (1.32 – 1.56) 1.03 (0.48 – 1.36) Growing ADG, kg/d 1.07 (0.76 – 1.51) 1.12 (0.58 – 2.40) 1.30 (0.69 – 1.83) 1.26 (0.77 – 2.07) Table 2. Meta-analysis of finishing performance and carcass traits from steers fed high- versus medium-starch growing diets prior to finishing from 9 published studies Item1 High Starch Medium Starch SEM P-value Performance Initial BW, kg Final BW, kg ADG, kg/d DMI, kg/d Gain:Feed 362.5 527.7 1.50 8.72 0.172 359.2 525.5 1.50 8.80 0.171 Carcass characteristics HCW, kg 323.9 325.6 2 LM area, cm 82.10 82.28 Rib fat thickness, cm 1.27 1.28 KPH, % 2.70 2.69 Yield Grade 2.94 2.90 2 Marbling Score 426.7 435.7 1 HCW = hot carcass weight, KPH = kidney, pelvic and heart fat. 2 Marbling grid: Slight00=300, Small00=400, Modest00=500. 8.1 16.4 0.10 0.32 0.008 0.21 0.59 0.78 0.62 0.62 10.6 2.39 0.09 0.25 0.18 17.4 0.45 0.78 0.63 0.59 0.50 0.35 42 Table 3. Meta-analysis of finishing performance and carcass traits from steers fed high- versus low-starch growing diets prior to finishing from 7 published studies Item1 High Starch Low Starch SEM P-value Performance Initial BW, kg Final BW, kg ADG, kg/d DMI, kg/d Gain:Feed 361.5 532.8 1.76 10.19 0.174 356.5 535.1 1.67 10.29 0.162 Carcass characteristics HCW, kg 331.8 330.8 2 LM area, cm 77.53 77.95 Rib fat thickness, cm 1.22 1.21 KPH, % 2.42 2.44 Yield Grade 3.02 3.11 Marbling score2 443.1 433.6 1 HCW = hot carcass weight, KPH = kidney, pelvic and heart fat. 2 Marbling grid: Slight00=300, Small00=400, Modest00=500. 13.1 11.4 0.07 0.36 0.010 0.38 0.61 0.09 0.58 0.12 10.2 3.27 0.10 0.27 0.16 19.1 0.77 0.68 0.88 0.75 0.15 0.40 Table 4. Summary statistics of 29 trials used in meta-analysis to evaluate the relationship of stocker phase ADG and initial finishing BW with carcass characteristics Variable N Mean Minimum Maximum Trial initial BW, kg 79 240.4 186.0 278.0 Stocker ADG, kg/d 85 0.77 0.15 1.68 Initial finishing BW, kg 85 346.7 231.5 450.0 Hot carcass weight, kg 85 326.1 240.2 397.0 LM area, cm2 78 78.80 64.50 96.57 Rib fat thickness, cm 82 1.30 0.51 2.40 Kidney, pelvic and heart fat, % 56 2.16 1.62 3.47 Yield Grade 73 2.99 2.37 4.02 Marbling score1 85 417 266 535 1 Marbling grid: Slight00=300, Small00=400, Modest00=500. 43 Table 5. Regression coefficients (± SE) of carcass characteristics on stocker phase ADG (kg/d), initial finishing BW (kg), or hot carcass weight (kg) from a meta-analysis of 29 trials Independent variable R2 Intercept HCW1, kg 0.788 0.939 12th rib fat, cm 0.001 0.062 0.734 1.2719 ± 0.0741 1.1751 ± 0.2803 -0.0335 ± 0.2661 0.0106 ± 0.0643*ADG 0.0002 ± 0.0007*initial finishing BW 0.0040 ± 0.0001*HCW LMA, cm2 0.419 0.695 0.864 75.7086 ± 1.5518 63.5700 ± 3.9409 34.5479 ± 5.1707 4.9965 ± 1.5243*ADG 0.0450 ± 0.0118*initial finishing BW 0.1367 ± 0.0158*HCW KPH, % 0.341 0.296 0.143 1.9745 ± 0.1144 1.3788 ± 0.2402 1.4338 ± 0.4911 0.2705 ± 0.0977*ADG 0.0023 ± 0.0006*initial finishing BW 0.0023 ± 0.0015*HCW Yield Grade 0.011 0.075 0.278 2.9268 ± 0.0971 2.6095 ± 0.3618 2.0057 ± 0.4756 0.0073 ± 0.0983*ADG 0.0009 ± 0.0010*initial finishing BW 0.0028 ± 0.0014*HCW 1 310.80 ± 10.95 224.94 ± 23.39 Independent variables 27.4394 ± 9.8434*ADG 0.2931 ± 0.0623*initial finishing BW HCW = hot carcass weight; LMA = longissimus muscle area; KPH = kidney, pelvic and heart fat. 44 550 Marbling Score 500 450 400 350 300 250 0.00 0.20 0.40 0.60 0.80 1.00 1.20 1.40 1.60 1.80 Stocker ADG, kg/d Figure 1. Mixed model regression of marbling score (♦) on ADG during the stocker or backgrounding phase without and with inclusion of other independent variables in the model. Marbling grid: 300 = Slight00; 400 = Small00; 500 = Modest00. Marbling score (solid line) = 403 ± 12.1 + 9.53 ± 9.44*ADG (R2 = 0.048). Marbling score (dashed line) = 311 ± 44.0 + 21.25 ± 7.86*ADG + 69.79 ± 34.35*ribfat (R2 = 0.79). Marbling score (dash/dotted line) = 176.2 ± 55.78 + 29.36 ± 22.29*ADG – 0.216 ± 0.175*initial finishing BW + 0.604 ± 0.274*HCW + 69.80 ± 28.48*ribfat (R2 = 0.84). 45 550 Marbling Score 500 450 400 350 300 250 200 250 300 350 400 450 500 Initial Finishing BW, kg Figure 2. Mixed model regression of marbling score (♦) on initial finishing weight without and with inclusion of other independent variables in the model. Marbling grid: 300 = Slight00; 400 = Small00; 500 = Modest00. Marbling score (solid line) = 392 ± 23.06 + 0.052 ± 0.060*initial finishing BW (R2 = 0.034). Marbling score (dashed line) = 282 ± 47.39 + 0.143 ± 0.053*initial finishing BW + 65.08 ± 34.64*ribfat (R2 = 0.77). Marbling score (dash/dotted line) = 176.2 ± 55.78 + 29.36 ± 22.29*ADG – 0.216 ± 0.175*initial finishing BW + 0.604 ± 0.274*HCW + 69.80 ± 28.48*ribfat (R2 = 0.84). 46 550 Marbling Score 500 450 400 350 300 250 200 250 300 350 400 450 Hot Carcass Weight, kg Figure 3. Mixed model regression of marbling score (♦) on hot carcass weight without and with inclusion of other independent variables in the model. Marbling grid: 300 = Slight00; 400 = Small00; 500 = Modest00. Marbling score (solid line) = 215 ± 93.05 + 0.596 ± 0.290*HCW (R2 = 0.76). Marbling score (dashed line) = 156 ± 51.28 + 0.478 ± 0.160*HCW + 76.29 ± 22.82*ribfat (R2 = 0.84). Marbling score (dash/dotted line) = 176.2 ± 55.78 + 29.36 ± 22.29*ADG – 0.216 ± 0.175*initial finishing BW + 0.604 ± 0.274*HCW + 69.80 ± 28.48*ribfat (R2 = 0.84). 47 450 Hot Carcass Weight, kg 400 350 300 250 200 0 0.2 0.4 0.6 0.8 1 1.2 1.4 1.6 1.8 Stocker ADG, kg/d Figure 4. Mixed model regression of hot carcass weight (♦) on ADG during the stocker or backgrounding phase with inclusion of initial finishing BW and ribfat in the model. Hot carcass weight (solid line) = 311 ± 10.95 + 27.44 ± 9.84*ADG (R2 = 0.79). Hot carcass weight (dashed line) = 154.4 ± 18.83 – 17.60 ± 8.60*ADG + 0.36 ± 0.068*initial finishing BW + 49.91 ± 8.46*ribfat (R2 = 0.94). 48 450 Hot Carcass Weight, kg 400 350 300 250 200 200 250 300 350 400 450 500 Initial Feedlot Placement Weight, kg Figure 5. Mixed model regression of hot carcass weight (♦) on initial finishing BW with inclusion of ADG during the stocker or backgrounding phase and ribfat in the model. Hot carcass weight (solid line) = 224.9 ± 23.39 + 0.29 ± 0.062*initial finishing BW (R2 = 0.94). Hot carcass weight (dashed line) = 154.4 ± 18.83 – 17.60 ± 8.60*ADG + 0.36 ± 0.068*initial finishing BW + 49.91 ± 8.46*ribfat (R2 = 0.94). 49