H. Berends

Low-protein solid feed improves the utilization of milk replacer for protein gain in veal calves

This study was designed to quantify the contribution of low-protein solid feed (SF) intake, in addition to milk replacer, to protein and energy retention in veal calves. Because of potential interactions between milk replacer and SF, occurring at either the level of digestion or postabsorption, this contribution might differ from that in calves fed either SF or milk replacer alone. Forty-eight Holstein Friesian male calves, 55±0.3 kg of body weight (BW), were divided across 16 groups of 3 calves each. Groups were assigned randomly to 1 of 4 incremental levels of SF intake: 0, 9, 18, or 27 g of DM of SF/kg of BW(0.75) per day. The SF mixture consisted of 25% chopped wheat straw, 25% chopped corn silage, and 50% nonpelleted concentrate (on a DM basis). Each group was housed in a respiration chamber for quantification of energy and N balance at each of 2 BW: at 108±1.1 kg and at 164±1.6 kg. The milk replacer supply was 37.3g of DM/kg of BW(0.75) per day at 108 kg of BW and 40.7 g of DM/kg of BW(0.75) per day at 164 kg of BW, irrespective of SF intake. Within a chamber, each calf was housed in a metabolic cage to allow separate collection of feces and urine. Indirect calorimetry and N balance data were analyzed by using regression procedures with SF intake-related variables. Nitrogen excretion shifted from urine to feces with increasing SF intake. This indicates a higher gut entry rate of urea and may explain the improved N utilization through urea recycling, particularly at 164 kg of BW. At 108 kg of BW, the gross efficiency of N retention was 61% for calves without SF, and it increased with SF intake by 5.4%/g of DM of SF per day. At 164 kg of BW, this efficiency was 49% for calves without SF, and it increased by 9.9%/g of DM of SF per day. The incremental efficiency of energy retention, representing the increase in energy retained per kilojoule of extra digestible energy intake from SF, was 41% at 108 kg of BW and 54% at 164 kg of BW. Accordingly, the apparent total-tract digestibility of NDF increased with BW, from 46% at 108 kg of BW to 56% at 164 kg of BW. On average, 5.5% of gross energy from SF was released as CH(4) in veal calves, which is similar to reported values in cattle fed only SF. In conclusion, the provision of low-protein SF resulted in improved N utilization for protein gain, particularly toward the end of the fattening period. In heavy calves, recycling of urea originating from amino acids in milk replacer potentially contributes substantially to the N retention of veal calves fed SF.

Effects of early rumen development and solid feed composition on growth performance and abomasal health in veal calves

The experiment was designed to study the importance of early rumen development and of the composition of solid feed intake on growth performance and abomasal health in milk-fed veal calves. One hundred and six Holstein-Friesian male calves were included in the experiment, and studied during 2 successive 12-wk periods (period 1 and period 2). In a 2 × 2 factorial arrangement, effects of partially replacing milk replacer by solid feed during period 1 and partially replacing dry matter (DM) intake from maize silage and barley straw by concentrate during period 2 were tested. Solid feed during period 1 consisted of maize silage, barley straw, and concentrate (25:25:50 on a DM basis). Solid feed during period 2 consisted of maize silage and barley straw (50:50 ratio on DM basis) for the nonconcentrate groups, and maize silage, barley straw and concentrates (25:25:50 on a DM basis) for the concentrate groups. At the end of period 1 (n=16) and at the end of period 2 (n=90), parameters of animal performance, rumen development, rumen fermentation, ruminal drinking, and abomasal damage were examined. Partially replacing milk replacer by solid feed during period 1 resulted in early rumen development (ERD) at the end of period 1, characterized by increased rumen weight, and an increased epithelial and absorptive surface area. Both ERD and partially replacing roughage by concentrates in period 2 increased the rumen development score at the end of period 2. Although ERD calves consumed more solid feed and less milk replacer during period 1 and 2 than non-ERD calves, carcass weight gains at 25 wk were identical, and utilization of the solid feed provided appeared similar to that of milk replacer. Partially replacing roughage by concentrates in period 2 increased dressing percentage and warm carcass weight. Plaque formation at the rumen mucosa was unaffected by ERD or partially replacing roughage by concentrates and generally low in all calves. The prevalence of large scars in the abomasum in ERD calves was decreased compared with non-ERD calves. This may indicate that ERD provided protection against abomasal lesions. In conclusion, early compared with late rumen development improves feed utilization and may be beneficial for abomasal health.

Urea Recycling Contributes to Nitrogen Retention in Calves Fed Milk Replacer and Low-Protein Solid Feed

Urea recycling, with urea originating from catabolism of amino acids and hepatic detoxification of ammonia, is particularly relevant for ruminant animals, in which microbial protein contributes substantially to the metabolizable protein supply. However, the quantitative contribution of urea recycling to protein anabolism in calves during the transition from preruminants (milk-fed calves) to ruminants [solid feed (SF)-fed calves] is unknown. The aim of this study was to quantify urea recycling in milk-fed calves when provided with low-protein SF. Forty-eight calves [164 ± 1.6 kg body weight (BW)] were assigned to 1 of 4 SF levels [0, 9, 18, and 27 g of dry matter (DM) SF · kg BW(-0.75) · d⁻¹] provided in addition to an identical amount of milk replacer. Urea recycling was quantified after a 24-h intravenous infusion of [¹⁵N₂]urea by analyzing urea isotopomers in 68-h fecal and urinary collections. Real-time qPCR was used to measure gene expression levels of bovine urea transporter B (bUTB) and aquaglyceroporin-3 and aquaglyceroporin-7 in rumen wall tissues. For every incremental gram of DM SF intake (g DM · kg(0.75)), nitrogen intake increased by 0.70 g, and nitrogen retention increased by 0.55 g (P < 0.01). Of this increase in nitrogen retention, 19% could be directly explained by urea recycling. Additionally, part of the observed increase in nitrogen retention could be explained by the extra protein provided by the SF and likely by a greater efficiency of postabsorptive use of nitrogen for gain. Ruminal bUTB abundance increased (P < 0.01) with SF provision. Aquaglyceroporin-3 expression increased (P < 0.01) with SF intake, but aquaglyceroporin-7 expression did not. We conclude that in addition to the increase in digested nitrogen, urea recycling contributes to the observed increase in nitrogen retention with increasing SF intake in milk-fed calves. Furthermore, ruminal bUTB and aquaglyceroporin-3 expression are upregulated with SF intake, which might be associated with urea recycling.

Estimation of milk leakage into the rumen of milk-fed calves through an indirect and repeatable method

In milk-fed calves, quantification of the milk that enters the rumen (ruminal milk volume, RMV) because of malfunction of the esophageal groove reflex may explain part of the variability observed between animals in their growth performance. The RMV can directly be quantified by adding an indigestible marker to the diet and measuring its recovery in the rumen at slaughter, but this technique cannot be repeated in time in the same animal. The objective of the study was to evaluate three indirect methods for estimating RMV. The first method was based on the assumption that ruminal drinking delays and limits acetaminophen appearance in blood after ingestion of milk supplemented with acetaminophen. The second method was based on a negative linear relationship between RMV and urinary recovery of non-metabolizable monosaccharides (3-O-methylglucose, l-rhamnose and d-xylose) added to the milk, owing to rumen fermentation. In the third method, RMV was calculated as the difference between total milk intake and the increase in abomasal milk volume (AMV) at feeding, measured through ultrasonography shortly after feeding, or estimated from the mathematical extrapolation of AMV to feeding time, based on consecutive measurements. These methods were tested in three experiments where calves (n=22, 10 and 13) were bucket fed or partly tube fed (i.e. by inserting milk replacer into the rumen via a tube to mimic ruminal drinking). In addition, Co-EDTA and Cr-EDTA were used as an indigestible marker in one experiment to trace bucket-fed or tube-fed milk replacer, respectively, to measure RMV. The relationship between AMV measured by ultrasonography and AMV measured at slaughter improved when kinetics of AMV were extrapolated to the time of slaughter by mathematical modeling (error between predicted and measured AMV equaled 0.49 l). With this technique, RMV during feeding averaged 17% and 24% of intake in Experiments 2 and 3, respectively. Plasma acetaminophen kinetics and recovery of non-metabolizable monosaccharides in urine were partly associated with ruminal drinking, but these techniques are not considered quantitatively accurate without further information of rumen degradation and absorption. The recovery of indigestible marker measured at slaughter gave a quantitative estimate of RMV (2% in Experiment 3), but improper measurement of emptying rate of fluid from the rumen may lead to underestimation. In conclusion, measuring changes in AMV by ultrasonography, in response to milk feeding, was the most promising indirect method to quantify RMV in veal calves.

Fermentation in the Small Intestine Contributes Substantially to Intestinal Starch Disappearance in Calves

BACKGROUND The proportion of starch disappearing from the small intestinal lumen is generally lower in ruminants than in monogastric animals, and there are indications that the starch digestion capacity in ruminants is limited. OBJECTIVES Milk-fed calves were used to study the rate-limiting enzyme in starch hydrolysis and to quantify starch fermentation in ruminants. METHODS Forty male Holstein-Friesian calves were fed milk replacer containing either lactose (control) or 1 of 4 corn starch products. The following starch products differed in the enzyme ratios required for their complete hydrolysis to glucose: gelatinized starch [α-amylase and (iso)maltase], maltodextrin [(iso)maltase and α-amylase], maltodextrin with α-1,6-branching (isomaltase, maltase, and α-amylase), and maltose (maltase). In the adaptation period, calves were stepwise exposed to an increasing dose of the starch product for 14 wk to allow maximal adaptation of all enzyme systems involved. In the experimental period, apparent total tract and ileal starch product disappearance, total tract starch product fermentation, and α-amylase, maltase, and isomaltase activities were determined at 18% inclusion of the starch product. RESULTS Maltase and isomaltase activities in the brush border did not increase for any of the starch product treatments. Luminal α-amylase activity was lower in the proximal (3.9 ± 3.2 and 2.7 ± 1.7 U/mg Co for control and starch product calves, respectively) but greater in the distal small intestine of starch-fed calves than in control calves (0.0 ± 0.0 and 6.4 ± 1.5 U/mg Co for control and starch product calves, respectively; means ± SEs for control and means ± pooled SEMs for starch product treatments). Apparent ileal (61.6% ± 6.3%) and total tract (99.1% ± 0.4%) starch product disappearance did not differ between starch product treatments, suggesting that maltase activity limits starch digestion in ruminants. Total tract starch product fermentation averaged 414 ± 43 g/d, corresponding to 89% of intake, of which half was fermented before the terminal ileum, regardless of starch product treatment. CONCLUSION Fermentation, rather than enzymatic digestion, is the main reason for small intestinal starch disappearance in milk-fed calves.

Utilization of roughages and concentrates relative to that of milk replacer increases strongly with age in veal calves

We aimed to investigate the feeding values of milk replacer (MR), roughage, and concentrates for veal calves in a paired-gain setting, thus avoiding any prior assumptions in feeding values and major differences in nutrient intakes. One hundred sixty male Holstein-Friesian calves at 2 wk of age and 45 ± 0.2 kg of body weight (BW) were included in the experiment. Calves were allocated to pens (5 calves per pen) and pens were randomly assigned to 1 of 4 solid feed (SF) levels: SF1, SF2, SF3, or SF4, respectively, and to 1 of 2 roughage-to-concentrate (R:C) ratios: 20:80 or 50:50. An adaptation period from wk 1 to 10 preceded the experimental period (wk 11 to 27). Total dry matter (DM) intake from SF was targeted to reach 20, 100, 180, and 260 kg of DM for SF1 to SF4, respectively, during the 16-wk experimental period, and increased with preplanned, equal weekly increments. Roughage was composed of 50% corn silage and 50% chopped wheat straw based on DM. The quantity of MR provided was adjusted every 2 wk based on BW to achieve similar targeted rates of carcass gain across treatments. The reduction in MR provided (in kg of DM) to realize equal rates of carcass gain with inclusion of SF (in kg of DM) differed between the R:C ratio of 50:50 (0.41 kg of MR/kg of SF) and the R:C ratio of 20:80 (0.52 kg of MR/kg of SF). As carcass gain unintentionally increased with SF intake, the paired-gain objective was not fully achieved. When adjusted for realized rates of carcass gain, calves fed an R:C ratio of 20:80 still required 10% less MR than calves fed an R:C ratio of 50:50 for equal rates of carcass gain, indicating that the utilization of SF for gain increased with concentrate inclusion. Averaged for the 16-wk experimental period, the feeding value of MR relative to that of concentrates and roughages was close to that predicted based on their respective digestible energy contents. Nevertheless, the feeding value of SF relative to that of MR increased substantially with age. Therefore, additivity in feeding values of these ration components cannot be assumed. The results of the current study may contribute to the development of new concepts for formulation of veal calf diets with substantial amounts of SF.

The role of solid feed amount and composition and of milk replacer supply in veal calf welfare

Diets used in veal production were linked with welfare problems: that is, behavioral and gastrointestinal health disorders. This study aimed to determine how indicators of calf welfare, that is, behavior and some characteristics of the feces reflecting gastrointestinal health status, are affected by (1) different amounts and compositions of solid feed (SF), (2) the addition of ad libitum long straw to a typical veal diet, and (3) milk replacer (MR) being fed via automated milk dispensers (AMD). Two-week-old Holstein-Friesian bull calves (n=270) were used in this study. In a 4×2 factorial design, 32 pens (5 calves per pen) were allocated to different levels of SF (SF1, SF2, SF3, or SF4) and roughage-to-concentrate ratios (20:80 or 50:50). The experimental period (13 to 29wk of age) was preceded by an adaptation period (3 to 12wk of age). Targeted total dry matter (DM) intake from SF during the experimental period was 20 kg of DM for SF1, 100 kg of DM for SF2, 180 kg of DM for SF3, and 260 kg of DM for SF4. The roughage part of the SF was 50% maize silage and 50% chopped wheat straw (on a DM basis). Ten additional pens were allocated to 2 treatments with ad libitum SF, with either (1) SF components in separate troughs (SEP) or (2) SF components mixed, with the composition being equal to the choice of SEP calves in the preceding week (MIX). Another 4 pens were fitted with racks filled with long wheat straw (STR) and fed SF2 with a roughage-to-concentrate ratio of 20:80. All the aforementioned pens received MR in buckets. Finally, 8 pens were allocated to 1 of 2 SF levels: SF1 or SF2 (with a roughage-to-concentrate ratio of 50:50) and fed MR via an AMD. Milk replacer provision was adjusted every 2wk to achieve similar rates of carcass gain across treatments (excluding SEP, MIX, and STR). Behavior was recorded at 15 and 24wk using instantaneous scan sampling. The prevalence of diarrhea and clay-like feces, which signal ruminal drinking, was monitored at 14 and 25wk. More roughage, but not concentrate, increased rumination and decreased tongue playing. The STR calves had higher rumination and lower abnormal behavior levels compared with calves without ad libitum straw. Offering MR via an AMD reduced tongue playing at 15wk. Tongue playing frequency was related to both roughage amount and AMD feeding, suggesting 2 separate motivations (i.e., rumination and sucking) underlying the development of this behavior. Only SF amount affected aspects of feces: SF1 calves had the highest diarrhea incidence. No effect of diet was found on clay-like feces.

A titration approach to identify the capacity for starch digestion in milk-fed calves

Calf milk replacers (MR) commonly contain 40% to 50% lactose. For economic reasons, starch is of interest as a lactose replacer. Compared with lactose, starch digestion is generally low in calves. It is, however, unknown which enzyme limits the rate of starch digestion. The objectives were to determine which enzyme limits starch digestion and to assess the maximum capacity for starch digestion in milk-fed calves. A within-animal titration study was performed, where lactose was exchanged stepwise for one of four starch products (SP). The four corn-based SP differed in size and branching, therefore requiring different ratios of starch-degrading enzymes for their complete hydrolysis to glucose: gelatinised starch (α-amylase and (iso)maltase); maltodextrin ((iso)maltase and α-amylase); maltodextrin with α-1,6-branching (isomaltase, maltase and α-amylase) and maltose (maltase). When exceeding the animals capacity to enzymatically hydrolyse starch, fermentation occurs, leading to a reduced faecal dry matter (DM) content and pH. Forty calves (13 weeks of age) were assigned to either a lactose control diet or one of four titration strategies (n=8 per treatment), each testing the stepwise exchange of lactose for one SP. Dietary inclusion of each SP was increased weekly by 3% at the expense of lactose and faecal samples were collected from the rectum weekly to determine DM content and pH. The increase in SP inclusion was stopped when faecal DM content dropped below 10.6% (i.e. 75% of the average initial faecal DM content) for 3 consecutive weeks. For control calves, faecal DM content and pH did not change over time. For 87% of the SP-fed calves, faecal DM and pH decreased already at low inclusion levels, and linear regression provided a better fit of the data (faecal DM content or pH v. time) than non-linear regression. For all SP treatments, faecal DM content and pH decreased in time (P<0.001) and slopes for faecal DM content and pH in time differed from CON; P<0.001 for all SP), but did not differ between SP treatments. Faecal DM content of SP-fed calves decreased by 0.57% and faecal pH by 0.32 per week. In conclusion, faecal DM content and pH sensitively respond to incremental inclusion of SP in calf MR, independently of SP characteristics. All SP require maltase to achieve complete hydrolysis to glucose. We therefore suggest that maltase activity limits starch digestion and that fermentation may contribute substantially to total tract starch disappearance in milk-fed calves.

Economic and environmental effects of providing increased amounts of solid feed to veal calves

Traditionally, veal calves receive most of their nutrients from milk replacer (MR). Nowadays, however, solid feed (SF; i.e., concentrates and roughages) increasingly substitutes for MR. Studies have shown that providing SF reduces different types of nonnutritive oral behaviors. The objective of this study was to assess the economic and environmental effects of substituting SF for MR in veal calf diets. With respect to environmental effects, we considered the emission of greenhouse gases and land occupation. Substitution rates were based on an experiment in which 160 calves were provided 2 mixtures of SF at 4 levels of dry matter (DM) intake. Mixtures of SF contained either 80% concentrates, 10% corn silage, and 10% straw on DM basis (C80) or 50% concentrates, 25% corn silage, and 25% straw (C50). The 4 levels of SF during the last 17 wk of the fattening period were 20, 100, 180, and 260 kg of DM SF. Additionally, provision of MR was adjusted to achieve equal rates of carcass gain. Substitution rates, representing the SF equivalent needed to substitute for 1 kg of DM MR, were 1.43 kg of DM for C80 and 1.61 kg of DM for C50. Economic effects were assessed based on prices and substitution rates of SF for MR and the possible penalty for carcass color. Environmental effects were assessed based on effects related to the production of feed ingredients, substitution rates, and changes in enteric methane emission and energy use for feed preparation. Costs of feeding SF needed to substitute for 1 kg of DM MR were €0.68 lower for C80 and €0.71 lower for C50, compared with the costs of feeding 1 kg of DM MR. When carcass color scores became too high, however, lower feeding costs were offset by lower revenues from meat. Emissions of greenhouse gases were hardly affected when SF intake was increased. In general, increased enteric methane emission were offset by lower emissions from feed production and energy use. Land occupation increased when intake of SF was increased, mostly because of the high land occupation associated with some concentrate ingredients. In conclusion, this study only showed a negative effect on land occupation when substituting SF for part of the MR in diets of veal calves. Effects on costs and greenhouse gas emissions were neutral or positive.

Effect of protein provision via milk replacer or solid feed on protein metabolism in veal calves

The current study evaluated the effects of protein provision to calves fed a combination of solid feed (SF) and milk replacer (MR) at equal total N intake on urea recycling and N retention. Nitrogen balance traits and [(15)N2]urea kinetics were measured in 30 calves (23 wk of age, 180±3.7kg of body weight), after being exposed to the following experimental treatments for 11 wk: a low level of SF with a low N content (SF providing 12% of total N intake), a high level of SF with a low N content (SF providing 22% of total N intake), or a high level of SF with a high N content (SF providing 36% of total N intake). The SF mixture consisted of 50% concentrates, 25% corn silage, and 25% straw on a dry matter basis. Total N intake was equalized to 1.8g of N·kg of BW(-0.75)·d(-1) by adjusting N intake via MR. All calves were housed individually on metabolic cages to allow for quantification of a N balance of calves for 5 d, and for the assessment of urea recycling from [(15)N2]urea kinetics. Increasing low-N SF intake at equal total N intake resulted in a shift from urinary to fecal N excretion but did not affect protein retention (0.71g of N·kg of BW(-0.75)·d(-1)). Increasing low-N SF intake increased urea recycling but urea reused for anabolism remained unaffected. Total-tract neutral detergent fiber digestibility decreased (-9%) with increasing low-N SF intake, indicating reduced rumen fermentation. Increasing the N content of SF at equal total N intake resulted in decreased urea production, excretion, and return to ornithine cycle, and increased protein retention by 17%. This increase was likely related to an effect of energy availability on protein retention due to an increase in total-tract neutral detergent fiber digestion (>10%) and due to an increased energy supply via the MR. In conclusion, increasing low-N SF intake at the expense of N intake from MR, did not affect protein retention efficiency in calves. Increasing the N content of SF at equal total N intake decreased urea production, increased protein retention, and coincided with improved fiber degradation. Therefore, results suggest that low N availability in the rumen limits microbial growth and rumen fermentation in calves fed low-N SF (93g of CP/kg of DM), and this effect cannot be compensated for by recycling of urea originating from MR.