Responses to incremental nutrient supply on energy and protein metabolism in pre-weaning dairy calves.

Recently reviewed development objectives and feeding practices in young dairy calves require an adaptation of nutrient recommendations set for milk replacer (MR) composition. Nutrient requirements of calves younger than 21 d of age, and those of calves fed with high levels of milk replacer are insufficiently quantified. The efficiency at which macronutrients are utilized, particularly protein, substantially diminishes with age, and there is little data for the first weeks of life. In addition, in older (pre-)ruminants, protein and energy can be simultaneously limiting for protein gain. Whether this also applies to calves in the first weeks of life is unknown. Therefore, this study aimed to quantify the responses in protein and fat gain to incremental supply of protein, fat, or lactose to MR in very young calves. Thirty-two groups of 3 mixed-sex Holstein-Frisian newborn calves (3.4 ± 1.6 d of age), were randomly assigned to one of 4 dietary treatments applied for 19 d: a basal MR (23.3% CP, 21.2% CF and 48.8% lactose of DM), provided at 550 kJ/kg BW0.85 per day (CON; n = 24), or the basal MR incrementally supplied with 126 kJ of DE/BW0.85 per day as milk fat (+FAT; n = 23), lactose (+LAC; n = 24) or milk protein (+PRO; n = 23). Calves were fed MR in 2 daily meals and had ad libitum access to water, but did not have access to calf starter nor any other solid feed. After 2 weeks of adaptation to their respective diets, groups of calves were placed for one week in an open-circuit respiration chamber for nitrogen and energy balance measurements (5 d). The incremental nutrient efficiencies indicate what percentage of extra intake of nutrients is retained. In this study, we observed that with every 100 g increase in protein intake, 52% was converted into protein deposition, while 44% contributed to heat production. Similarly, a 100 g increase in fat intake resulted in 67% being stored as fat, 22% being released as heat, and only 5% being retained as protein. Likewise, a 100 g increment in lactose intake led to 49% being stored as fat, with 38% being released as heat. Additional protein intake was not deposited as fat, extra energy intake (fat, and additional lactose) increased post absorptive N efficiency in young calves.

Effects of mixing a high-fat extruded pellet with a dairy calf starter on performance, feed intake, and digestibility.

During weaning, withdrawal of milk replacer is not directly compensated for by an increase in solid feed intake. Therefore, greater fat inclusion in the starter might mitigate this temporary dietary energy decline. However, fat inclusion in solid feeds may generally limit rumen fermentability and development. To address these potentially conflicting outcomes, we conducted 2 experiments to evaluate the effect of supplementing a high-fat extruded pellet mixed with a calf starter on feed intake, performance, and nutrient digestibility in calves. In experiment 1, 60 Holstein bull calves were blocked by serum IgG (2,449 ± 176 mg/dL) and date of arrival (2.5 ± 0.5 d of age). Within each block, calves were randomly assigned to 1 of 3 treatments: a standard control calf starter (CON; 3.1% fat) and mixtures of CON with 10% inclusion of 1 of 2 different high-fat extruded pellets containing 85% of either hydrogenated free palm fatty acids (PFA, 7.1% fat) or hydrogenated rapeseed triglycerides (RFT, 6.7% fat). Calves were offered milk replacer up to 920 g/d until 42 d of age, followed by a gradual weaning period of 7 d. Calves had ad libitum access to the starter diets, straw, and water. No differences were observed between CON, PFA, and RFT calves on body weight (BW) or average daily gain (ADG) until 49 d of age. From weaning (50 d) until 112 d, PFA calves had a greater BW and ADG than RFT and CON animals. Moreover, PFA calves had the highest intakes of starter, straw, calculated metabolizable energy, and crude protein after weaning. Overall, no differences were present in blood ß-hydroxybutyrate and glucose concentrations between treatments; however, calves in the RFT treatment had a higher concentration of insulin-like growth factor-1. In experiment 2, 24 Holstein bull calves at 3 mo of age were assigned to 1 of 8 blocks based on arrival BW and age. Within each block, calves were randomly assigned to 1 of the 3 treatments previously described for experiment 1. Calves on the RFT treatment had the lowest total-tract apparent dry matter and fat digestibility, potentially explaining the differences in performance observed between PFA and RFT calves. Inclusion of the PFA pellet at 10% with a calf starter improved BW, solid feed, and energy intake after weaning. However, these benefits were conditioned by fat source and its digestibility.

Metabolic and performance responses to the replacement of lactose by fat in milk replacer formulations for dairy calves.

The recent trend in the dairy industry towards elevated planes of milk feeding of young calves requires reconsideration of calf milk replacer (CMR) formulations. The fat:lactose ratio in CMR is typically lower than that of whole milk and effects of increasing fat inclusion at the expense of lactose in CMR on nutrient metabolism and gut function of rearing calves are not sufficiently understood. Therefore, the current study aimed to determine the effect of increasing replacement of lactose by fat on growth performance, nutrient digestibility and metabolism, and glucose/insulin sensitivity. A total of 40 male calves (1.7 ±â€¯0.10 days of age, 46.7 ±â€¯0.76 kg BW) were blocked based on arrival date and randomly assigned to one of four treatments containing differing levels of fat and lactose (18F: 18.8% and 47.6%; 22F: 22.3% and 42.8%; 26F: 26.0% and 38.6%; 30F: 30.1% and 33.8%, fat and lactose DM, respectively). Calves were individually housed for the duration of the 11 week study and received their CMR (150 g/L) twice a day (0700 and 1600 h) from a teat bucket. The CMR feeding schedule consisted of 6 L/d from d 2 to 14, 7 L/d from d 15 to 56, and then 4 L/d during gradual weaning from 56 to 63 days. Pelleted starter, chopped straw, and water were available ad libitum throughout the study. Measurements included feed intake, growth, nutrient digestibility, fecal composition, and blood parameters. A glucose tolerance test (GTT) was performed between 28 and 32 days of age. By design, metabolizable energy intake from CMR increased linearly with fat level, but this did not result in a difference in BW, ADG, or concentrate intake. Fecal composition remained unaffected by treatment except for higher fat content in 22F compared to 26F. Also, plasma non-esterified fatty acids and total bilirubin differences were limited to 22F having the highest values whereas 26F had the lowest values. Regarding the GTT, total area under the curve (AUC) for glucose was highest in 22F and 26F and lowest in 18F whereas the AUC between 30 and 60 min for glucose was highest in 26F and lowest in 18F and 22F. Overall, altering the lactose:fat ratio in CMR did not affect growth performance while having minor effects on nutrient metabolism, but future investigation should focus on physiological consequences and effects in adult life to understand biological impact of fat and lactose levels in CMR for rearing calves.

Determining the nutritional boundaries for replacing lactose with glucose in milk replacers for calves fed twice daily.

The effect of replacing lactose with glucose on the gastrointestinal system of young calves at levels above 20% diet inclusion in milk replacer (MR) is not well described. The aim of this study was to determine tolerance to glucose inclusion at the direct expense of lactose on glucose metabolism, health, and growth performance in Holstein male calves. In total, 110 Holstein male dairy calves (16 ± 2.5 d and 50.3 ± 0.2 kg) were acquired from a commercial collection center. After an adaptation period of 3 d, 100 calves were selected for the study based on health parameters. Calves were blocked based on body weight measured on d 4 after arrival. Within each block, calves were randomly assigned to 1 of 5 levels of glucose inclusion (replacing lactose): 0% (L1, n = 20), 10% (L2, n = 20), 20% (L3, n = 20), 30% (L4, n = 20), and 40% (L5, n = 20), leading to an estimated osmolality range from 417 (L1) to 586 mOsm/kg (L5). Carbohydrates were exchanged based on hexose equivalents, and glucose delivery was standardized across treatments, while the rest of the formula (60%) remained unchanged. Calves received L1 during the adaptation period of 3 d and were then exposed to their respective treatment until d 47 after arrival. Milk replacer was provided daily in 2 equally sized meals. Meal size was 2.0 L during the 3-d adaptation period and gradually increased to 4.0 L until weaning (d 35 after arrival). During weaning, meal size decreased from 4.0 to 2.0 L on d 36, and MR was withdrawn on d 48 after arrival. Straw and concentrates were offered ad libitum from d 25 onward. Calves had ad libitum access to water throughout the study. Measurements included daily feed intakes, weekly body weight, and weekly spot feces sampling in all calves. Blood samples were collected on d 18. Additionally, postprandial responses of insulin and glucose were measured in 6 calves per treatment on d 19, 20, and 21. Increasing glucose inclusion (at the direct expense of lactose) in MR did not affect growth but linearly increased mortality, which was as high as 25% (5/20) in L5. Mortality was primarily associated with gastrointestinal disorders (6/11). At higher glucose levels, calves needed greater serum insulin concentrations to control glycemia, as shown by a linear increase in the area under the curve for insulin. Furthermore, calves needed more time to control glycemia, as indicated by a linear increase in the maximal concentration of insulin. Consequently, there was a linear increase in area under the curve for glucose. Even though calves needed more time and higher insulin concentrations for 30% glucose inclusion and higher, the glucose-to-insulin ratio did not differ across treatments. However, high glucose inclusion levels in MR affected calf mortality and is not a suitable strategy for lactose replacement.

Effects of exchanging lactose for fat in milk replacer on ad libitum feed intake and growth performance in dairy calves.

The recent trend in the dairy industry toward ad libitum feeding of young calves merits reconsideration of calf milk replacer (CMR) formulations. Additionally, feed intake regulation in young calves provided with ad libitum milk and solid feeds is insufficiently understood. This study was designed to determine the effect of exchanging lactose for fat in CMR on voluntary feed intake and growth performance. Lactose was exchanged for fat on a weight/weight basis, resulting in different energy contents per kilogram of CMR. Thirty-two male calves (1.7 ± 0.12 d of age, 47.6 ± 0.83 kg of body weight) were assigned to 1 of 16 blocks based on arrival date. Within each block, calves were randomly assigned to 1 of 2 treatments. The experimental period was divided into 4 periods. In period 1, until 14 ± 1.7 d of age, calves were individually housed, restricted-fed their assigned CMR treatments at 2.5 to 3 L twice daily, and provided with unlimited access to water, chopped straw, and starter. In period 2, calves were group-housed with 8 calves per pen and received ad libitum access to their assigned CMR treatments, starter feed, chopped wheat straw, and water. During period 3, from 43 until 63 d of age, calves were weaned by restricting CMR allowance in 2 steps, maintaining access to all other feeds. All calves were completely weaned at d 64 of age and were monitored until 77 d of age (period 4). Measurements included the intake of all dietary components, body weight gain, and a selection of blood traits. Increasing fat content at the expense of lactose decreased CMR intake by 10%, whereas total calculated metabolizable energy intake and growth remained equal between treatments. Total solid feed (starter and straw) consumption was not affected by CMR composition. These data indicate that calves fed ad libitum regulate their CMR intake based on energy content. High-fat CMR increased plasma phosphate, nonesterified fatty acids, triglycerides, and bilirubin, whereas plasma glucose remained unchanged. Despite the limited animal numbers in the present experiment, there was a significant decrease in the total number of health events (mainly respiratory) requiring therapeutic intervention and in the total number of therapeutic interventions in calves fed high-fat CMR. Calves appeared to consume CMR based on energy content, with a difference in ad libitum intake proportional to the difference in energy content of the CMR, maintaining equal body weight gain and solid feed intake.

Effect of energy source in calf milk replacer on performance, digestibility, and gut permeability in rearing calves.

Current calf milk replacer (CMR) compositions significantly differ from whole milk in their levels of energy, protein, and minerals. Energy source is one of the major differences, as CMR contains high levels of lactose, whereas whole milk contains higher levels of fat. The aim of this study was to determine the effect of partially exchanging lactose for fat on performance, digestibility, and gut permeability in calves fed twice daily on a high feeding plane. Lactose and fat were exchanged in the CMR formulation on a weight-weight basis. The CMR were isonitrogenous but not isoenergetic. A total of 60 male Holstein-Friesian calves were assigned to 1 of 30 blocks based on serum IgG, body weight, and date of collection after birth. Within each block, calves were randomly assigned to 1 of 2 treatments: high fat and high lactose. The CMR was provided twice daily until 49 d of age, followed by a gradual weaning period of 14 d. Starter, straw, and water were available ad libitum throughout the complete study. Exchanging lactose for fat did not affect growth; intakes of starter, straw, water, crude protein, or total energy; or apparent total-tract digestibility of nutrients. Gastrointestinal permeability was assessed by measuring the recovery of lactulose and Cr in 24-h urine and the Cr concentration and lactulose:d-mannitol ratio in serum following an oral pulse dose. Urinary recoveries of Cr and lactulose were generally low in both treatments but were higher in calves fed the high-fat CMR. Accordingly, the serum lactulose:d-mannitol ratio and serum Cr concentrations were higher in calves fed the high-fat CMR. In wk 1 and during the weaning transition, calves fed the high-fat CMR had significantly fewer abnormal fecal scores. In conclusion, exchanging lactose for fat in the CMR did not affect growth performance, total feed intake, or nutrient digestibility. The high-fat CMR was associated with an increase in permeability markers but positively influenced fecal scores in calves.

Preweaning nutrient supply alters mammary gland transcriptome expression relating to morphology, lipid accumulation, DNA synthesis, and RNA expression in Holstein heifer calves.

The objective of this study was to analyze the mammary gland transcriptome to determine how preweaning nutrient supply alters the molecular mechanisms that regulate preweaning mammary development. Holstein heifers were fed via milk replacer (MR) either an elevated level of nutrient intake (ELE; on average, 5.9 ± 0.2 Mcal of ME in 8.4 L of MR/d, n = 6) or a restricted amount of nutrients (RES; 2.8 ± 0.2 Mcal of ME in 4 L of MR/d, n = 5) for 54 d after birth, at which point they were slaughtered and samples of mammary parenchyma tissue were obtained. Parenchymal mRNA was analyzed, and the fold change (FC) of 18,111 genes (ELE relative to RES) was uploaded to Ingenuity Pathway Analysis (IPA) software (Qiagen Bioinformatics, Redwood City, CA) for transcriptomic analysis. Using a threshold of P < 0.05, IPA identified that the FC of 1,931 of 18,811 differentially expressed genes (DEG) could be used for the analysis. A total of 18 molecular and cellular functions were relevant to DEG arising from the treatments; the 5 functions most associated with DEG were cell death and survival, cellular movement, cellular development, cellular growth and proliferation, and lipid metabolism. Based on the directional FC of DEG, the mammary gland of ELE heifers was predicted to have increased epithelial-mesenchymal transition (Z = 2.685) and accumulation of lipid (Z = 2.322), whereas the synthesis of DNA (Z = -2.137), transactivation of RNA (Z = -2.254), expression of RNA (Z = -2.405), transcription (Z = -2.482), and transactivation (Z = -2.611) were all predicted to be decreased. Additionally, IPA predicted the activation status of 13 upstream regulators with direct influence on DEG as affected by ELE feeding that were ligand-dependent nuclear receptors (n = 2), enzymes (n = 1), or transcription regulators (n = 10). Of these, 6 were activated (Z > 2) and 7 were inhibited (Z < -2). In summary, feeding ELE preweaning altered the mammary transcriptome of Holstein heifers, affecting cell functions involved in the morphological and physiological development of the mammary gland.

Effects of fat inclusion in starter feeds for dairy calves by mixing increasing levels of a high-fat extruded pellet with a conventional highly fermentable pellet.

The objective of this study was to evaluate the effect of inclusion of an extruded high-fat pellet mixed with a conventional pelleted calf starter on energy intake and performance around weaning in calves. To this end, 75 female Holstein newborn calves (41.0 ± 4.98 kg) were randomly assigned to 1 of 5 iso-nitrogenous solid feed treatments consisting of 4 levels of fat inclusion by mixing a low-fat highly fermentable control pellet with 3 different levels of inclusion of an extruded high-fat pellet [control (100:0), 90:10, 80:20, and 70:30], and a high-fat single pellet (HFSP). The HFSP was equivalent iso-energetic and iso-nitrogenous, although it had almost 1 percentage point difference in fat relative to the 80:20 treatment, to contrast the effect of the dual-component pellet mixture. The extruded high-fat starter feed contained a high proportion of fat (38%), mainly from hydrogenated palm fatty acids. Calves were offered a milk replacer up to 900 g/d, and then pre-weaned at 49 d of age by halving milk allowance until 56 d when calves were weaned. Calves had ad libitum access to the starter diets, chopped straw, and water. Individual milk replacer and starter intakes were recorded daily and BW was determined weekly. A glucose tolerance test was performed at 49 and 84 d of age to evaluate blood glucose homeostasis. Apparent total-tract digestibility was determined from 70 to 75 d of age. Calves on the 90:10 treatment had greatest starter feed intake mainly due to a marked increase in solid feed intake around weaning. Metabolizable energy intake was increased when the extruded pellet was included in the starter. No differences were present in digestibility of ether extract among solid feed treatments. The area under the curve of blood glucose concentration after the glucose tolerance test was greatest in 80:20; intermediate in 70:30, HFSP, and control; and lowest in 90:10 calves. No differences were observed for insulin or other parameters related to blood glucose homeostasis. Delivering dietary fat by mixing an extruded high-fat pellet with a conventional highly fermentable pellet to reach a total fat content of 7% results in increased starter intake, energy intake, and body weight gain until 84 d of age compared with a conventional low-fat pellet, or a single pellet with increased fat content.

Short communication: Parameters of abomasal emptying and glucose-insulin dynamics in Holstein-Friesian calves at 2 ages and 2 levels of milk replacer intake.

Elevated planes of nutrition in the preweaning period of dairy calf management can increase average daily gain, shorten age at puberty, and increase milk yield. In a previous study, 12 Holstein calves were fed 2 meals/d of 4 or 2 L milk replacer up to 7 wk of age. The objective of the current study was to estimate parameters of abomasal emptying and glucose-insulin dynamics in these calves by fitting a mechanistic model to postprandial appearances of plasma glucose, insulin, and the abomasal emptying marker acetaminophen measured at 4 and 7 wk of age. Higher intake of milk replacer resulted in longer bouts of abomasal emptying at a slower rate. Parameters of glucose and insulin dynamics were not affected by milk replacer intake. However, older calves had decreased insulin-stimulated glucose utilization indicating impaired insulin sensitivity, as well as increased pancreatic responsiveness. Neither of these effects were apparent from i.v. glucose tolerance tests on the calves and may have been related to postprandial gut hormone release. Effects of age on parameters of glucose-insulin dynamics were larger than effects of milk replacer intake. Conversely, effects of milk replacer intake on abomasal emptying were larger than effects of age.

Understanding oral stereotypies in calves: alternative strategies, hypothalamic-pituitary-adrenal axis (re)activity and gene by environment interactions.

Stereotypies are used as indicators of poor animal welfare and it is, therefore, important to understand underlying factors mediating their development. In calves, two oral stereotypies, that is, tongue playing and object manipulation, result mostly from insufficient structure in the diet. Three hypotheses were studied: (1) oral stereotypies in calves are one of two alternative strategies, the alternative being hypo-activity; (2) stereotyping and non-stereotyping calves differ in terms of cortisol secretion; (3) oral stereotypy development in calves rests on a gene by environment interaction. Eight-week-old bull calves (n=48) were assigned to one of four solid feed allowances (0, 9, 18 or 27 g dry matter/kg metabolic weight per day) with the following composition: 50% concentrate, 25% maize silage and 25% straw on dry matter basis. The calves received milk replacer in buckets, the provision of which was adjusted to achieve equal growth rates. At 14 to 18 weeks of age, calves were exposed to a challenge, that is, tethering inside cages. Oral stereotypies and inactivity were recorded in the home pens in the 4 weeks before the challenge using instantaneous scan sampling. Salivary cortisol levels were measured at -120, +40, +80, +120 min and +48 h relative to the challenge. Individual differences in behaviour were recorded in the first 30 min after challenge implementation using focal animal sampling and continuous recording, and these elements were entered into a principal component (PC) analysis to extract PCs. Regression analyses were performed to find relationships between stereotypies and inactivity, stereotypies and cortisol, and stereotypies and PCs (individual differences, genes) and solid feed (environment). Relationships between PCs and cortisol were also investigated to help with the interpretation of PCs. Hypotheses 1 and 2 were rejected. Hypothesis 3, however, was supported: calves with a zero solid feed allowance, that is, in the most barren environment, showed links between stereotypies and two of the PCs. Calves that displayed high levels of idle and rapid locomotion and low levels of oral contact with the cage during the challenge also displayed high levels of object manipulation in the home pens. Calves that displayed low levels of stepping and turning attempts during the challenge also displayed high levels of tongue playing in the home pens. This study corroborates the gene by environment interaction on the development of oral stereotypies in calves.

Effect of plane of milk replacer intake and age on glucose and insulin kinetics and abomasal emptying in female Holstein Friesian dairy calves fed twice daily.

The objective of this study was to investigate how preweaning plane of milk replacer intake and age can affect insulin and glucose kinetics as well as abomasal emptying rate in dairy calves fed twice a day. A total of 12 female Holstein Friesian calves were blocked by cow parity, paired by colostrum origin, and were randomly assigned to a high plane of milk replacer intake (8 L/d, 1.2kg of milk replacer/d; n=6) or a low plane of nutrition (4 L/d, 0.6kg of milk replacer/d; n=6). All calves received 4 L of colostrum over 2 meals (1 and 6h after birth) and were then directly transferred to their assigned feeding plans until they were stepped-down from milk by 50% during wk 7 and weaned on wk 8. Milk replacer (24% crude protein, 18% crude fat) was fed at 150g/L twice daily (0700 and 1700h) and all calves had ad libitum access to pelleted calf starter, chopped wheat straw, and water. Jugular catheters were placed in all calves at 4, 7, and 10wk of age. Then, postprandial response to plasma glucose, insulin, and acetaminophen (supplied with the meal) were determined to measure abomasal emptying. The next day, a glucose tolerance test was conducted by infusing glucose via the jugular catheter. At 4 and 7wk of age, the rate constant (%/h) for abomasal emptying of the meal was lower in high calves (0.21±0.02 in wk 4; 0.27±0.02 in wk 7) compared with low (0.34±0.02 in wk 4; 0.47±0.02 in wk 7). The postprandial plasma insulin area under the curve over 420min was greater in high calves (18,443±7,329; low=5,834±739 µU/mL) compared with low. We found no differences in glucose tolerance test kinetics between the high and low dairy calves at 4, 7, or 10wk of age. The findings from this study suggest that feeding dairy calves an elevated plane of nutrition in 2 meals of milk replacer per day does not decrease insulin sensitivity.

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.

Effects of solid feed level and roughage-to-concentrate ratio on ruminal drinking and passage kinetics of milk replacer, concentrates, and roughage in veal calves.

Effects of solid feed (SF) level and roughage-to-concentrate (R:C) ratio on ruminal drinking and passage kinetics of milk replacer, concentrate, and roughage were studied in veal calves. In total, 80 male Holstein-Friesian calves (45±0.2kg of body weight) were divided over 16 pens (5 calves per pen). Pens were randomly assigned to either a low (LSF) or a high (HSF) SF level and to 1 of 2 R:C ratios: 20:80 or 50:50 on a dry matter (DM) basis. Roughage was composed of 50% corn silage and 50% chopped wheat straw on a DM basis. At 27 wk of age, measurements were conducted in 32 calves. During the measurement period, SF intake was 1.2kg of DM/d for LSF and 3.0kg of DM/d for HSF, and milk replacer intake averaged 2.3kg of DM/d for LSF and 1.3kg of DM/d for HSF. To estimate passage kinetics of milk replacer, concentrate, and straw, indigestible markers (CoEDTA, hexatriacontane C36, Cr-neutral detergent fiber) were supplied with the feed as a single dose 4, 24, and 48h before assessment of their quantitative recovery in the rumen, abomasum, small intestine, and large intestine. Rumen Co recovery averaged 20% of the last milk replacer meal. Recoveries of Co remained largely unaffected by SF level and R:C ratio. The R:C ratio did not affect rumen recovery of C36 or Cr. Rumen fractional passage rate of concentrate was estimated from recovery of C36 in the rumen and increased from 3.3%/h for LSF to 4.9%/h for HSF. Rumen fractional passage rate of straw was estimated from Cr recovery in the rumen and increased from 1.3%/h for LSF to 1.7%/h for HSF. An increase in SF level was accompanied by an increase in fresh and dry rumen contents. In HSF calves, pH decreased and VFA concentrations increased with increasing concentrate proportion, indicating increased fermentation. The ratio between Cr and C36 was similar in the small and large intestine, indicating that passage of concentrate and straw is mainly determined by rumen and abomasum emptying. In conclusion, increasing SF level introduces large variation in passage kinetics of dietary components, predominantly in the rumen compartment. The SF level, rather than the R:C ratio, influences rumen recovery of concentrate and roughage. Our data provide insight in passage kinetics of milk (Co representing the milk replacer) and SF (Cr and C36 representing roughage and concentrate, respectively) and may contribute to the development of feed evaluation models for calves fed milk and 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 animal's 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.

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.

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.

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.

Evaluation of the SF6 tracer technique for estimating methane emission rates with reference to dairy cows using a mechanistic model.

A dynamic, mechanistic model of the sulfur hexafluoride (SF6) tracer technique, used for estimating methane (CH4) emission rates from ruminants, was constructed to evaluate the accuracy of the technique. The model consists of six state variables and six zero-pools representing the quantities of SF6 and CH4 in rumen and hindgut fluid, in rumen and hindgut headspace, and in blood and collection canister. The model simulates flows of CH4 and SF6 through the body, subsequent eructation and exhalation and accumulation in a collection canister. The model predicts CH4 emission by multiplying the SF6 release rate of a permeation device in the rumen by the ratio of CH4:SF6 in collected air. This prediction is compared with the actual CH4 production rate, assumed to be continuous and used as a driving variable in the model. A sensitivity analysis was conducted to evaluate the effect of changes in several parameters. The predicted CH4 emission appeared sensitive to parameters affected by the difference in CH4:SF6 ratio in exhaled and eructed air respectively, viz., hindgut fractional passage rate and hindgut CH4 production. This is caused by the difference in solubility of CH4 and SF6 and by hindgut CH4 production. In addition, the predicted CH4 emission rate appeared sensitive to factors that affect proportions of exhaled and eructed air sampled, i.e., eructation time fraction, exhalation time fraction, and distance from sampling point to mouth/nostrils. Changes in rumen fractional passage rate, eructation rate, SF6 release rate, background values and air sampling rate did not noticeably affect the predicted CH4 emission. Simulations with (13)CH4 as an alternative tracer show that the differences and sensitivity to parameters greatly disappear. The model is considered a useful tool to evaluate critical points in the SF6 technique. Data from in vivo experiments are needed to further evaluate model simulations.

Differential cortical activation during observation and observation-and-imagination.

The activity of the brain during observation or imagination of movements might facilitate the relearning of motor functions after stroke. The present study examines whether there is an additional effect of imagination over observation-only. Eight healthy subjects observed and observed-and-imagined a movement of a hand; 64-channel EEG was used to measure brain activity. The synchronization of the theta (4-8 Hz), alpha (8-13 Hz) and beta (13-25 Hz) frequency bands was calculated and plotted in topoplots. The temporal changes of the sensorimotor area (C3, C4) and the centro-parietal cortex (Pz) were analyzed in the two experimental conditions. During observation-and-imagination, a significant larger desynchronization (p = 0.004) in the sensorimotor area was found compared to observation-only in all electrodes and frequency bands. In addition, temporal differences were found between observation and observation-and-imagination in the alpha frequency bands. During observation-and-imagination, modulations of EEG rhythms were stronger than during observation-only in the theta, alpha and beta frequency bands and during almost the whole activity fragment. These findings suggest an additive effect of imagination to observation in the rehabilitation after stroke.