1. Introduction
In beef meat production systems, carcass and meat traits depend on breed, age, and diets of slaughtered animals. Previous studies have reported that the tenderness of meat has a negative correlation with the age of cull cows [
1,
2] due to the chemical nature of connective tissue [
3]. Furthermore, negative attributes associated with carcass and meat from cull cows, including lower carcass weights, smaller
Longissimus dorsi muscle (LM) areas, inferior muscling, yellower external fat color, and darker lean color [
4,
5]. Generally, providing high-energy ration to cull cows before slaughter improves carcass characteristics and meat quality [
6,
7] and reverses such an influence.
The inclusion and the type of forage in the diet of finishing cattle depend upon the availability, price, and its influence on fattening and meat quality [
8]. Providing forage in sufficient amounts is considered as the key factor in preventing ruminal acidosis due to increased saliva production [
9]. However, excessive amounts of forage tend to reduce dry matter intake (DMI), which may be related to its big bulk, slow rumen passage rate and low apparent digestibility [
10,
11] and consequently influence finishing performance. Therefore, the appropriate amounts of forage are required to maintain rumen health and maximize finishing performance.
In high marbled beef production system, especially with cull cows, the study on the effect of age and forage levels in the diet could be of great significance. In previous studies with cull cows, few experiments have been conducted dealing with the effect of the breed [
1,
12], finishing diet [
13,
14], and age [
1,
2]. They have shown that breed and interaction with finishing diet can modify the fat deposition in the carcass and meat, the muscle fiber type, the collagen content and composition, and in consequence the meat quality in terms of tenderness, flavor, and juiciness. Furthermore, few experiments have studied the relationships of different types of finishing diets with beef meat quality of heifers [
15], young bulls [
16], and cull beef cows [
17], but they did not study factors such as age and their interaction with finishing diet.
Yiling cattle originated in Yichang City and has been identified as a new breed of Chinese yellow cattle. Its history as a draft animal can be traced back to 4000 years. Yiling cattle is currently considered as a draught and meat type breed. Previous studies have evaluated the genetic background of Yiling cattle [
18], but the information about the finishing performance and meat traits of Yiling cattle is still partially unclear.
Antioxidant systems are an important determinant of the oxidative stability of stored meat, influenced by breed, age, diet, and environment. The aging process is caused by deleterious effects of reactive oxygen species (ROS) generated spontaneously from normal cellular metabolism [
19]. In beef production, cull cows had higher myoglobin or heme-Fe contents [
20], which accelerates the oxidation process [
21]. Previous study elaborated that inclusion of forage in the diet of beef cattle during the finishing phase altered antioxidant status [
22]. However, the antioxidative status of tissues (i.e., liver and muscle tissues) in cull cows in the finishing phase concerning animal age is still needed to be investigated.
For these reasons, our objectives were to assess the effects of age and rice straw inclusion levels in the diet of Yiling cattle on carcass characteristics, and intramuscular fat (IMF) content, tenderness and water-holding capacity of meat. In the current study, we hypothesized that the reduction of the amounts of straw in the finishing diet of cows would show better beef quality due to the potential enhancement of energy intake. Besides, younger cull cows may have greater finishing performance due to their higher eating appetite and capacity. The other objective of the current study was to evaluate the tissue antioxidant status of Yiling cull cows. We hypothesized that older age would induce protein and lipid oxidation of tissues and consequently enhance its antioxidant capacity. Moreover, dietary forage levels may alter antioxidant status of cull cows under high-energy ration conditions.
4. Discussion
One of the most important parameters of beef production is the growth ability of animals. Growth in young cattle is unequal and influenced by age and diet. In the current study, younger cows showed higher final BW, total BW gain, and average daily gain, which is supported by the study of Sawyer et al. (2004) [
28], who found that DMI and ADG decreased linearly with the increased age of the cull cows. However, the results in growth performance are opposite to the finding of Galli et al. (2008) [
29], who reported that younger cows finished under grazing conditions had lower the final BW than older cows. The increased DMI with forage inclusion levels in the diet could be supported by Galyean and Defoor (2003) [
30]. They reported a positive linear relationship (
R2 = 0.92) between forage NDF and DMI (% BW), and also claimed that rumen and gut fill does not limit intake when ruminants are fed high concentrate diets and the mechanism for enhanced intake is energy dilution and to maintain energy intake. However, we believe the changed intake is more likely due to the more balanced ruminal environment and the greater ruminal pH value, particularly in high concentrate diet, caused by enhanced chewing activity and saliva flow with increasing forage inclusion [
7]. In the present study, calculated by the proportions of total concentrate and straw intake in total DMI during finishing period, the percentages of forage in diet groups were 15.2% and 21.7% (LRS vs. HRS), respectively. Hales et al. (2013) [
31] compared different levels of forage inclusion (2%, 6%, 10%, or 14% of alfalfa hay) and showed a quadratic effect of increasing forage proportion on DMI during the whole finishing period in steers. They further reported that an increase in DMI with alfalfa hay inclusion of up to 10%, and then a decrease in DMI with 14% alfalfa hay. A previous study by Swanson et al. (2017) [
32] reported that ADG and DMI decreased linearly with increasing forage inclusion. The inconsistency among studies may be due to the differences in breed or forage type. The apparent digestibility of OM (slightly) and NDF decreased with straw levels, which is consistent with the result of Salinas-Chavira et al. (2013) [
33]. Generally, Chinese south native cattle were used as draft animals and had not undergone long-term commercial selection. Thus, we speculated that Yiling cattle may be intolerant to forage restriction diet that could cause intake inhibition.
In terms of feed intake, daily gain and feed conversion efficiency, this fattening strategy seems less efficient in resource utilization. The inefficiency may be mainly due to the fact that the experimental cattle were mature cows and the increase in BW comes from fat deposition rather than the growth and development of bones and muscles. Although there is no control group (no fattening) in the current study, the production experience of the farm has proved that long-term fattening can increase the marbling richness of beef from Yiling cull cows. High marbled beef, especially from native cattle, can fetch high prices at the market, which can support the reasonableness of this fattening strategy.
Younger cull cows and cows fed higher straw levels had heavier carcass that could be justified by their higher final BW. A recent study reported that the proportion of fat in carcasses increased, and the relative proportion of muscles decreased as animals become older [
34]. However, fat percentage in carcass was similar between age groups in the current study. Dressing percentage is an important indicator in the evaluation of carcass characteristics, and the rise in dressing percentage is a direct result of increasing fatness with slaughter weight [
35]. Thus, similar dressing percentage between experimental groups was due to unchanged fat contents in carcass in the current study. In addition, heavier striploin found in cows fed with higher straw levels and high rib found in younger cows were related to higher carcass weight. Taken together, the present results may not only be useful for further studies, but also for the sustainable and profitable beef production from cull cows.
In the current study, no significant significance, including the effects of age and diet and their interaction, was observed for IMF deposition. However, the mean value of IMF content in younger cows fed higher straw levels was 41.92% higher than in younger cows fed lower straw levels (31.0% vs. 21.9%). By contrast, the numerical difference in IMF content of older cows between diet groups was small (28.8% vs. 28.5%), which may indicate that IMF deposition is insensitive to the change of energy intake for older cows. It is known that the tenderness of beef from cull cows decreased with increasing age [
1,
2]. However, the shear force was unaffected by age in this study. A recent study [
36] reported that the shear force of beef from cull cows was lower than that from heifers when fed with a high-energy ration for 150 d, which indicates that the high IMF content induced by long-term fattening could weaken the effect of connective tissue properties on tenderness. Drip loss, pressing loss as well as cooking loss can describe the water-holding capacity of beef and reflect different characteristics. Galli et al. (2008) [
29] reported that cooking loss was unchanged with the increasing age of cull cows, which is consistent with our results. However, another recent study [
37] showed that age altered the cooking loss of beef from cull cows and this influence depends on different beef cuts. In the current study, dietary treatment has no effect on the physical properties of beef except for cooking loss. The decreased cooking loss caused by high-level straw diet may lead to inferior juiciness.
Fatty acid composition can vary dramatically in beef depending on several factors, such as breed, sex, age, and diet [
38]. Fruet et al. (2018) [
39] reported that higher C18:1n9c and MUFAs proportion as well as lower C20:5n3c and SFAs proportion were found in the high concentrate diet treatment. Moreover, Wang et al. (2019) [
27] reported that the proportions of C18:1n9c and MUFAs rose with dietary energy. In the current study, younger cows fed higher straw levels had higher C18:1n9c and MUFAs proportion as well as lower C20:5n3c and SFAs proportion than younger cows fed lower straw levels, which could be associated with the enhancement of energy intake. Although total TDN intake of two age classes both increased with straw levels, the effects of diet on the proportions of C15:0, C17:0, C18:1n9c, C20:5n3c, SFAs, and MUFAs were more pronounced in younger cows than in older cows. Several studies have demonstrated that the fatty acid profile was strongly affected by IMF content [
40,
41]. Thus, the interaction between age and diet for those fatty acids may partially come from their numerical difference in IMF content. Cho et al. (2013) [
42] reported that the proportions of C18:3n3c, C18:3n6c, SFA, and n-3 PUFAs increased and MUFAs decreased in striploin as the cow age increased. However, in this study, differences were only detected for the slightly higher n-6/n-3 PUFAs ratio and greater C18:3n3c proportion in older cows. Reasons for the inconsistency in these results may be related to the differences in age classes.
Oxidative stress caused by the increased production of ROS or a decrease in antioxidant capacity, results in damage to biological macromolecules as well as disruption of normal metabolism and physiology [
43]. Moreover, oxidative stress plays a key role in the pathogenesis of diverse diseases in cattle [
44,
45]. Antioxidases CAT, SOD, and glutathione peroxidase (GSH-Px) are part of enzymatic antioxidant systems that protect tissue components against oxidative stress caused by ROS. The concentration of MDA reflects the extent of lipid oxidation, which has a negative impact on meat freshness and quality, including undesirable off-flavor, toxic substances, and discoloration [
46]. Protein carbonyl (PC) is a biomarker of protein oxidative damage, which could decrease the water-holding capacity and tenderness of the meat [
47]. Halliwell (1994) [
48] reported that animal age induced the oxidative damage to cellular macromolecules, such as lipids, proteins, and DNA. Moreover, Cho et al. (2015) [
20] reported that oxidative deterioration on d2 post-slaughter was accelerated with older age, despite the increased activity of antioxidant enzymes. Enhanced activities of antioxidant enzymes during aging were the consequences of increased expression of the mRNA of antioxidant enzymes [
49]. However, only the CAT of liver tissue was different between age groups in this study, and its activities were higher in younger cows than in older cows. In addition, the antioxidant parameters were unaffected by straw amounts except for SOD. Antioxidase CAT and SOD are coupled enzymes [
50], but the activity of CAT and SOD in both liver and muscle tissues did not exhibit the same pattern in our results, most likely due to the additional effect of dietary stress in our study, including high energy intake and abnormal level of dietary forage. A high concentration of unsaturated fatty acids, particularly PUFAs, accelerates the lipid oxidation process [
51]. In the current study, the unsaturation degree in meat was unaffected by animal age and straw amounts, which may be related to the similarity of MDA concentration in muscle between experimental groups. The results that T-AOC, ROS, PC and MDA remain unchanged indicate that the antioxidant status under high-concentration diets may be less affected by age and diet.