S. L. Hansen

Effects of increased dietary sulfur on beef steer mineral status, performance, and meat fatty acid composition

Ninety-six crossbred yearling steers (321 ± 29 kg BW) were used to determine the effects of feeding cattle a high S diet on pasture before receiving a high S diet in the feedlot. Steers were blocked by BW, allocated to 2.4-ha bromegrass (Bromus inermis L.) pastures (n = 4 plots per treatment), and supplemented at 1% BW with either low S dried distillers grains with solubles (DDGS; 0.34% total diet S; LS) or LS DDGS with additional S (0.47% total diet S; HS) from NaSO(4) for 36 d. On d 37, steers moved into the feedlot where one-half remained on the previous S treatment and the other half switched treatments, resulting in 4 treatments (LS-LS, LS-HS, HS-LS, HS-HS; LS: 0.2 to 0.3% total diet S, HS: 0.5 to 0.6% total diet S; n = 6 feedlot pens per treatment). During the pasture period, forage mass offered, grazing residual mass, and in vitro digestible DM of forage did not differ among treatments (P > 0.40), and ADG did not differ (LS: 1.6 kg · d(-1), HS: 1.7 kg · d(-1), P = 0.54). Plasma Mg measured on d 35 was decreased by ≈ 5% in response to increased dietary S during the pasture period (P = 0.05), though no effect on plasma Mg was observed during finishing (P > 0.15). Plasma Cu concentrations on d 155 were ≈ 15% less (P = 0.02) in HS vs. LS steers, and d 155 liver Cu concentrations were ≈ 51% less in HS vs. LS steers (P = 0.01). Increased dietary S during the feedlot period decreased ADG by ≈ 10% (P = 0.01) and tended to decrease HCW by ≈ 5% (P = 0.06) compared with LS steers. Steers receiving the HS diet had increased stearic acid (C18:0) and heptadecanoic acid (C17:0; P = 0.04 and 0.01, respectively) percentages in rib facings collected at slaughter. Exposing cattle to greater S diets (0.47% S) during a forage-based diet did not influence later performance on high S feedlot diets (0.5 to 0.6% S); however, cattle fed high dietary S on pasture had greater fat cover at slaughter (P = 0.01), suggesting S may have influenced lipid metabolism.

Iron Transporters Are Differentially Regulated by Dietary Iron, and Modifications Are Associated with Changes in Manganese Metabolism in Young Pigs

To investigate the effects of dietary iron (Fe) on manganese (Mn) metabolism, 24 weaned pigs (21 d old) were blocked by litter and weight and randomly assigned to the following treatments: 1) no supplemental Fe [low Fe (L-Fe)]; 2) 100 mg supplemental Fe/kg [adequate Fe (A-Fe)]; and 3) 500 mg supplemental Fe/kg [high Fe (H-Fe)]. The basal diet was analyzed to contain 20 mg Fe/kg. Tissues were harvested after 32 d of feeding. Daily gain (least square means +/- SEM) was greater in A-Fe pigs (328.3 +/- 29.9 g/d) than in L-Fe pigs (224.0 +/- 11.2 g/d). Hemoglobin concentrations on d 32 were lower in L-Fe pigs (62 +/- 3.5 g/L) than in A-Fe pigs (128 +/- 5.6 g/L) and did not differ between pigs fed A-Fe and H-Fe (133 +/- 12.0 g/L). Liver Fe increased with increasing dietary Fe. Relative hepatic hepcidin expression was greater in pigs fed A-Fe and H-Fe than in those fed L-Fe. Relative expressions of duodenal divalent metal transporter 1 (DMT1) and solute carrier family 39 member 14 (ZIP14) were increased in L-Fe pigs compared with H-Fe pigs. Liver copper (Cu) was higher in L-Fe (0.56 +/- 0.04 mmol/kg) and H-Fe (0.58 +/- 0.04 mmol/kg) pigs than in A-Fe pigs (0.40 +/- 0.04 mmol/kg). Liver Mn was lower in H-Fe pigs (0.15 +/- 0.01 mmol/kg) than in A-Fe (0.23 +/- 0.02 mmol/kg) or L-Fe pigs (0.20 +/- 0.02 mmol/kg). Duodenal Mn concentrations were greater in L-Fe pigs than in A-Fe or H-Fe pigs. Fe deficiency in pigs increased gene expression of duodenal metal transporters (DMT1 and ZIP14) and supplementation with H-Fe reduced expression of DMT1 and ZIP14, which may have decreased absorption of Mn.

High dietary iron reduces transporters involved in iron and manganese metabolism and increases intestinal permeability in calves

A 56-d experiment was designed to examine the effect of high dietary Fe on metal transporters involved in Fe and Mn metabolism. Fourteen weaned Holstein calves were stratified by weight and randomly assigned to 1 of 2 treatments: 1) no supplemental Fe (normal Fe) or 2) 750mg of supplemental Fe/kg of dry matter (high Fe). Jugular blood was collected on d 0, 35, and 56. At the end of the trial, 6 calves per treatment were humanely killed and duodenal scrapings, liver, and heart were collected for analysis. Additionally, proximal duodenum was mounted on Ussing chambers to assess intestinal barrier integrity. Calves receiving high dietary Fe displayed decreased transepithelial resistance and increased apical-to-basolateral flux of radiolabeled mannitol, suggesting that high Fe created increased intestinal permeability. Feeding calves a diet high in Fe decreased average daily gain, dry matter intake, and feed efficiency. Hemoglobin and serum Fe concentrations did not differ due to dietary treatment. High dietary Fe increased concentrations of Fe in the liver, but did not affect heart or duodenal Fe concentrations. Duodenal Mn concentrations were lowered by feeding a high Fe diet, but liver and heart Mn concentrations were not affected. As determined by real-time reverse transcription PCR, relative hepatic expression of the gene that encodes the Fe regulatory hormone hepcidin was 5-fold greater in calves fed high dietary Fe. Hepcidin is released in response to increased Fe status and binds to the Fe export protein ferroportin causing ferroportin to be degraded, thereby reducing dietary Fe absorption. Confirmation of this result was achieved through Western blotting of duodenal protein, which revealed that ferroportin was decreased in calves fed high dietary Fe. Duodenal protein expression of divalent metal transporter 1 (DMT1), a Fe import protein that can also transport Mn, tended to be reduced by high dietary Fe. Transcript levels of several genes involved in Fe metabolism in liver and duodenum were unchanged by treatment. In summary, feeding calves a diet high in Fe induced a signal cascade (hepcidin) designed to reduce absorption of Fe (via reduced protein expression of ferroportin and DMT1) in a manner similar to that reported in rodents. Additionally, reduced levels of DMT1 protein appeared to decrease duodenal Mn, suggesting that Mn may also be a substrate for DMT1 in cattle.

Supplemental vitamin C improves marbling in feedlot cattle consuming high sulfur diets

The objective of this study was to examine the effects of supplemental rumen-protected vitamin C (VC) on live and carcass-based performance, and antioxidant capacity of cattle consuming varying concentrations of dietary S. Angus-cross steers (n = 120) were blocked by initial BW (341 ± 11 kg) and assigned equally to 1 of 6 treatments, evaluating 3 concentrations of dietary S [0.22%, 0.34%, and 0.55%, for low S (LS), medium S (MS), and high S (HS), respectively] and 2 concentrations of supplemental VC (0 or 10 g • steer(-1) • d(-1)). Steers receiving VC-supplemented diets consumed an average of 10.3 g of supplemental VC • steer(-1) • d(-1) and increasing dietary S linearly increased (P < 0.01) grams of S consumed. Increasing dietary S decreased (P < 0.01) DMI, final BW, and ADG, and linearly increased (P < 0.05) rumen hydrogen sulfide and blood sulfhemoglobin concentrations. The inclusion of VC, regardless of S treatment, tended to increase (P = 0.08) plasma VC concentrations, specifically within the medium and high S diets (P = 0.04). Plasma total antioxidant capacity (d 90) linearly decreased (P = 0.003) and total liver glutathione (GSH; d 143) tended to decrease (P = 0.08) due to increased S intake. Within the high S treatment, addition of VC decreased (P = 0.04) the ratio of oxidized-to-reduced GSH compared with HS alone. Increased dietary S and VC decreased (P < 0.05) plasma Cu concentrations, whereas VC increased (P = 0.01) plasma Fe concentrations. Linear decreases (P < 0.02) in marbling score, backfat thickness (BF), yield grade, and HCW were observed as dietary S increased; however, the addition of VC to the HS diet increased (P < 0.01) BF, marbling scores, and percentage of cattle grading Choice compared with HS without VC. In conclusion, supplementation of VC to cattle receiving the high S diet improved marbling scores; although the exact mechanism for this improvement is unknown, it may be related to greater circulating VC available for lipid metabolism in these cattle.

Bioaccessibility of iron from soil is increased by silage fermentation

High dietary Fe can negatively affect absorption of other minerals and cause tissue damage through the production of free radicals. Cattle are often exposed to high dietary Fe, and soil ingestion may represent a major dietary source of Fe. Iron in soil is often found in the ferric form bound in insoluble complexes; however, exposure to an acidic environment similar to that occurring during silage fermentation may cause this Fe to be reduced to the more soluble ferrous form. To test this theory, a 2 x 2 x 3 factorial arrangement examining time, level, and type of soil addition to greenchop was used. Factors included 2 times of soil addition (before or after ensiling), 2 levels of soil inclusion (1 and 5% contamination, wet basis) and 3 types of soil (Cecil clay loam, 3.4% Fe; Georgeville silt loam, 4.3% Fe; and Dyke clay loam, 6.9% Fe). In addition, greenchop with no soil added was ensiled to serve as a control. Fresh corn greenchop was mixed with the appropriate type and level of soil and tightly packed in experimental silos. Fermentation was allowed to proceed for 90 d before silos were opened and silage was freeze-dried and ground. To simulate contamination after ensiling, each soil type was added to control silage at the 2 levels of inclusion. Addition of soil to greenchop before ensiling resulted in greater amounts of water soluble Fe compared with soil addition after ensiling, suggesting that Fe-soil binding properties were altered by ensiling. To test the potential bioaccessibility of Fe during ruminant digestion, an enzymatic in vitro system was modified to simulate ruminal, abomasal, and intestinal digestion. The presence of soil, regardless of time of addition, type, or inclusion level, resulted in greater soluble or bioaccessible Fe concentrations after all 3 phases when compared with control silage. Ensiling further increased soluble Fe concentrations after each phase when compared with silage contaminated with soil after ensiling. In addition, dialyzable Fe concentration (15,000 Da molecular weight cut off) following intestinal phase simulation was greater due to ensiling. Iron that becomes soluble during the intestinal phase may be available to the animal for absorption, and ensiling resulted in increased concentrations of potentially bioavailable Fe. These results suggest that soil contamination of harvested feeds before ensiling may represent a major source of bioavailable Fe in the diets of cattle.

High-sulfur in beef cattle diets: A review

While many cattle feeding areas in the United States have long dealt with high sulfate water, increased feeding of ethanol coproducts such as distillers grains with solubles to beef cattle has led to a corresponding increase in dietary sulfur. As a result, sulfur metabolism in the ruminant has been the focus of many research studies over the past 10 yr, and advances in our knowledge have been made. Excessive sulfur in cattle diets may have implications on trace mineral absorption, dry matter intake, and overall cattle growth. This review will focus on what we have learned about the metabolism of sulfur in the ruminant, including ruminal sulfate reducing bacteria, the role of ruminally available sulfur, factors affecting the production of hydrogen sulfide in the rumen, and the potential mechanisms behind sulfur toxicity in cattle. Additionally, this review will discuss potential strategies to minimize risk of sulfur toxicity when cattle are fed high-sulfur diets, including dietary and management strategies. Further research related to high-sulfur diets including implications for carcass characteristics, meat quality, and animal health will also be discussed. As ethanol production processes continue to change, the nutrient profile of the resulting coproducts will as well. Often removal of one nutrient such as oil will result in the concentration of other nutrients such as sulfur. Therefore, it seems even more likely that a better understanding of sulfur metabolism in the ruminant will be important to beef cattle feeding in the future.

The addition of high manganese to a copper-deficient diet further depresses copper status and growth of cattle

A study was conducted evaluating the effect of long-term Cu deficiency, with or without high Mn, on growth, gene expression and Cu status of beef cattle. Twenty-one Angus calves were born to cows receiving one of the following treatments: (1) 10 mg supplemental Cu/kg DM (+Cu); (2) no supplemental Cu and 2 mg Mo/kg DM ( - Cu); (3) - Cu diet plus 500 mg supplemental Mn/kg DM ( - Cu+Mn). Calves were weaned at approximately 183 d of age and individually fed throughout the growing and finishing phases. Plasma Cu was lower (P < 0.01) in - Cu calves compared with +Cu calves while high dietary Mn further depressed (P < 0.01) plasma Cu in - Cu+Mn calves v. - Cu calves. Liver Cu concentrations in +Cu calves were greater (P < 0.01) than in - Cu calves, with no differences between - Cu and - Cu+Mn calves. The daily body-weight gain of +Cu calves was greater (P < 0.01) than - Cu calves during the period from birth to weaning, but did not differ during the growing phase. - Cu+Mn calves gained less (P < 0.05) than - Cu calves during the growing phase. DM intake was lower (P < 0.01) in - Cu+Mn calves v. - Cu calves, and did not differ among +Cu and - Cu calves. The relative gene expression of cytochrome c oxidase in the liver was lower (P < 0.05) in - Cu calves compared with +Cu or - Cu+Mn calves. In conclusion, feeding a Cu - deficient diet in combination with high Mn negatively affected the growth and Cu status of beef cattle.

Assessment of ruminal hydrogen sulfide or urine thiosulfate as diagnostic tools for sulfur induced polioencephalomalacia in cattle

To determine if ruminal hydrogen sulfide, urine thiosulfate, or blood sulfhemoglobin could be used as diagnostic indicators for sulfur-induced polioencephalomalacia, 16 steers (8 cannulated, 368 ± 12 kg; 8 unmodified, 388 ± 10 kg; mean ± standard error) were fed 1 of 2 dietary treatments. Diets consisted of a low sulfate (0.24% S; control) wheat midd–based pellet or the control pellet with sodium sulfate added to achieve a high-sulfate (0.68% S) pellet. As designed, intake did not differ (P = 0.80) between treatments. At 8 hr postfeeding, ruminal hydrogen sulfide was not affected by cannulation (P = 0.35) but was greater (P < 0.01) in high S (6,005 ± 475 mg/l) than control (1,639 ± 472 mg/l) steers. Time of day of sampling affected (P = 0.01) ruminal hydrogen sulfide, with peak concentrations occurring 4–12 hr after feeding. Urine was collected prefeeding (AM) and 7–9 hr postfeeding (PM). Urine thiosulfate concentrations of high S steers sampled in the PM were greater (P > 0.01) than in the AM. However, there was no difference due to time of sampling for control. In both the AM and PM, urine thiosulfate concentrations of high S were greater (P > 0.01) than control. Although hydrogen sulfide and thiosulfate were elevated by increased dietary S intake, a concentration at which polioencephalomalacia is likely to occur could not be determined. Sampling urine for thiosulfate or rumen gas for hydrogen sulfide of nonsymptomatic pen mates 4–8 hr after feeding may be useful to assess sulfur exposure and differentiate between causes of polioencephalomalacia.

High dietary sulfur decreases the retention of copper, manganese, and zinc in steers

To examine the effects of dietary S on diet digestibility and apparent mineral absorption and retention, 16 steers [8 ruminally fistulated (368 ± 12 kg BW) and 8 unmodified (388 ± 10 kg BW)] were paired within modification status and BW, and within each of the 2 consecutive 28-d periods, 4 pairs of steers were randomly assigned to either a low-S (0.24%) or high-S (0.68%) pelleted diet. Bromegrass hay was fed at 5 or 7% of the diet, during periods 1 and 2, respectively. Sodium sulfate was used to increase the S content of the high-S diet. The low-S steers were fed the amount of feed their high-S counterpart consumed the previous day, while the high-S steers received 110% of the previous days intake. Steers were adapted to individual metabolism stalls for 4 d (d -3 to 0 of period), acclimated to diet for 7 d (d 1 to 7 of period), and after high-S steers were consuming ad libitum intake for 7 d (d 14 of period), total urine and feces were collected for 5 d. Feed intake and orts were recorded daily. Dry matter and OM digestibility were determined. Jugular blood was collected before and after each collection period on d 14 and 20, and liver biopsies were collected on d 0 and 27. Macromineral (Ca, K, Mg, and Na) and micromineral (Cu, Mn, and Zn) concentrations were determined for pellets and hay, orts, feces, urine, and plasma and liver samples from each steer via inductively coupled plasma spectrometry. Dry matter intake, DM and OM digestibility, and urine volume were not affected (P ≥ 0.11) by dietary treatment, but fecal output was greater (P = 0.02) in the low-S steers than the high-S steers. A high-S diet decreased plasma Cu (P = 0.04) and liver Zn (P = 0.03) compared to low-S steers. No differences (P ≥ 0.20) were noted among urinary excretion of Cu, Mn, and Zn. Sodium absorption was greater (P < 0.01) and Cu, Mn, and Zn retention was lesser (P ≤ 0.01) in the high-S steers than the low-S steers. Apparent absorption of Ca, K, and Mg was not affected (P ≥ 0.18) by dietary treatment, while absorption of Cu, Mn, and Zn in the high-S treatment was lesser (P ≤ 0.06). In conclusion, consumption of a high-S diet for 28 d had limited effects on Ca, K, Mg, and Na absorption and retention, but decreased Cu, Mn, and Zn retention, which may limit growth and production of cattle consuming a high-S diet long-term.

Technical note: Copper chaperone for copper, zinc superoxide dismutase: A potential biomarker for copper status in cattle

Copper chaperone for Cu, Zn superoxide dismutase (CCS) has been shown to be reflective of Cu status in mice and rats. The objective of this study was to evaluate liver and erythrocyte CCS as an indicator of Cu status in beef cattle (Exp. 1), and to test the acute-phase properties of CCS under conditions of inflammation (Exp. 2). In Exp. 1, samples of whole blood and liver were collected at slaughter (492 d of age) from 15 Cu-deficient and 6 Cu-adequate Angus calves. At the time of tissue collection, severe Cu deficiency had been achieved and differences (P < 0.0001) in plasma and liver Cu among Cu-adequate and Cu-deficient calves were extreme (1.26 vs. 0.19 mg/L and 208.4 vs. 6.3 mg/kg for plasma and liver Cu, respectively). Protein levels of CCS were greater in liver (40%; P = 0.02) and erythrocytes (65%; P < 0.0001) of Cu-deficient vs. Cu-adequate calves. In Exp. 2, inflammatory responses were elicited in beef heifers by administration of a Mannheimia hemolytica vaccine. Four days after vaccination, plasma concentrations of the Cu-dependent protein ceruloplasmin and the Cu-independent protein haptoglobin were increased (P < 0.001) by 71 and 83%, respectively. In contrast, detection of CCS protein in samples of liver and erythrocytes did not differ (P >or= 0.45) between baseline (d 0) and d 4 after vaccination. These data demonstrate that bovine erythrocyte and liver CCS protein levels increase in Cu-deficient cattle. Furthermore, levels of CCS protein do not change after a vaccine-induced inflammatory response, suggesting that unlike ceruloplasmin, CCS may be a reliable indicator of Cu status in cattle.