Hostname: page-component-78c5997874-dh8gc Total loading time: 0 Render date: 2024-11-05T04:56:24.346Z Has data issue: false hasContentIssue false

Review: Importance of colostrum supply and milk feeding intensity on gastrointestinal and systemic development in calves

Published online by Cambridge University Press:  06 February 2020

H. M. Hammon*
Affiliation:
Institute of Nutritional Physiology ‘Oskar Kellner’, Leibniz Institute for Farm Animal Biology (FBN), Wilhelm-Stahl-Allee 2, Dummerstorf 18196, Germany
W. Liermann
Affiliation:
Institute of Nutritional Physiology ‘Oskar Kellner’, Leibniz Institute for Farm Animal Biology (FBN), Wilhelm-Stahl-Allee 2, Dummerstorf 18196, Germany
D. Frieten
Affiliation:
Department of Life Sciences and Engineering, University of Applied Sciences Bingen, Berlinstrasse 109, Bingen am Rhein 55411, Germany
C. Koch
Affiliation:
Educational and Research Centre for Animal Husbandry, Hofgut Neumuehle, Münchweiler an der Alsenz 67728, Germany
*

Abstract

Feeding management of the postnatal and preweaning calf has an important impact on calf growth and development during this critical period and affects the health and well-being of the calves. After birth, an immediate and sufficient colostrum supply is a prerequisite for successful calf rearing. Colostrum provides high amounts of nutrient as well as non-nutrient factors that promote the immune system and intestinal maturation of the calf. The maturation and function of the neonatal intestine enable the calf to digest and absorb the nutrients provided by colostrum and milk. Therefore, colostrum intake supports the start of anabolic processes in several tissues, stimulating postnatal body growth and organ development. After the colostrum feeding period, an intensive milk feeding protocol, that is, at least 20% of BW milk intake/day, is required to realise the calf potential for growth and organ development during the preweaning period. Insufficient milk intake delays postnatal growth and may have detrimental effects on organ development, for example, the intestine and the mammary gland. The somatotropic axis as the main postnatal endocrine regulatory system for body growth is stimulated by the intake of high amounts of colostrum and milk and indicates the promotion of anabolic metabolism in calves. The development of the forestomach is an important issue during the preweaning period in calves, and forestomach maturation is best achieved by solid feed intake. Unfortunately, intensive milk-feeding programmes compromise solid feed intake during the first weeks of life. In the more natural situation for beef calves, when milk and solid feed intake occurs at the same time, calves benefit from the high milk intake as evidenced by enhanced body growth and organ maturation without impaired forestomach development during weaning. To realise an intensive milk-feeding programme, it is recommended that the weaning process should not start too early and that solid feed intake should be at a high extent despite intensive milk feeding. A feeding concept based on intensive milk feeding prevents hunger and abnormal behaviour of the calves and fits the principles of animal welfare during preweaning calf rearing. Studies on milk performance in dairy cows indicate that feeding management during early calf rearing influences lifetime performance. Therefore, an intensive milk-feeding programme affects immediate as well as long-term performance, probably by programming metabolic pathways during the preweaning period.

Type
Review Article
Copyright
© The Animal Consortium 2020

Implications

Successful calf rearing depends on an enhanced colostrum management and is improved by an intensive milk-feeding programme that allows calves to develop their potential for growth and organ maturation. Feeding regimes with a restricted milk supply of 4 to 6 kg milk or milk replacer/day focus on forestomach development and may disregard body growth and the maturation of other visceral organs beyond the rumen. An ad libitum milk-feeding programme stimulates calf growth and organ development and is consistent with calf well-being and avoids hunger. In addition, intensive milk feeding may have a strong impact on calf health and long-life performance in cattle.

Introduction

Mortality and morbidity rates are still unacceptably high during early calf rearing. The incidence for mortality in the perinatal period, defined as the duration from birth to 48 h after birth, ranges in dairy herds worldwide between 3% and 9% (Compton et al., Reference Compton, Heuer, Thomsen, Carpenter, Phyn and McDougall2017). A recent survey on mortality rates in Germany revealed up to 17% calf losses (calf losses after birth up to 6 months of age) in dairy farms (Tautenhahn, Reference Tautenhahn2017). In US dairy herds, current mortality rates of 5% and morbidity rates of 34% were published for preweaning calves (Urie et al., Reference Urie, Lombard, Shivley, Kopral, Adams, Earleywine, Olson and Garry2018). The UK Department of the Environment, Food and Rural Affairs reported that economic losses from calf mortality were around £60 million/year (DEFRA, 2003). It is obvious that the high mortality and morbidity rates contradict the aim of increased animal welfare for farm animals and compromise the breeding of robust animals (Huber, Reference Huber2018). In dairy farming, calves usually do not grow up with their dam, and calves are immediately separated from their dams after birth. Thus, farmers are highly responsible for the colostrum and milk feeding management and can significantly contribute to an improved calf health and the reduction of calf losses during the postnatal period.

Feeding management during the neonatal and preweaning period has a great impact on the success of calf rearing and, in addition, affects health and performance in later life (Khan et al., Reference Khan, Weary and von Keyserlingk2011; Ballou, Reference Ballou2012; Van Amburgh and Soberon, Reference Van Amburgh and Soberon2013). Because severe diarrhoea is a main reason for neonatal calf losses, the management of milk feeding and especially colostrum supply in the first days of life is of particular importance for the success of calf rearing (DEFRA, 2003; Tautenhahn, Reference Tautenhahn2017; Urie et al., Reference Urie, Lombard, Shivley, Kopral, Adams, Earleywine, Olson and Garry2018). An adequate and immediate (within 2 to 3 h after birth) colostrum supply is important for establishing passive immunity in calves, and the amount of colostrum fed to newborn calves directly correlates with the prevention of illness and calf losses (Godden, Reference Godden2008; Mee, Reference Mee2008).

Furthermore, there is increasing evidence that an enhanced milk or milk replacer (MR) feeding schedule during the preweaning period not only affects growth but also promotes organ development and well-being (Geiger et al., Reference Geiger, Parsons, James and Akers2016; Schäff et al., Reference Schäff, Gruse, Maciej, Mielenz, Wirthgen, Hoeflich, Schmicke, Pfuhl, Jawor, Stefaniak and Hammon2016; Rosenberger et al., Reference Rosenberger, Costa, Neave, von Keyserlingk and Weary2017). This review aims to summarise the research on the impact of colostrum supply and subsequent intensive milk feeding on the gastrointestinal and systemic development and maturation of the preweaning calf. An intensive milk feeding schedule orientates on a daily milk intake of 20% instead of 10% of BW (Khan et al., Reference Khan, Weary and von Keyserlingk2011), which is closely related to ad libitum milk (Jasper and Weary, Reference Jasper and Weary2002; Schiessler et al., Reference Schiessler, Nussbaum, Hammon and Blum2002; Maccari et al., Reference Maccari, Wiedemann, Kunz, Piechotta, Sanftleben and Kaske2015) or MR feeding (Schäff et al., Reference Schäff, Gruse, Maciej, Mielenz, Wirthgen, Hoeflich, Schmicke, Pfuhl, Jawor, Stefaniak and Hammon2016; Frieten et al., Reference Frieten, Gerbert, Koch, Dusel, Eder, Kanitz, Weitzel and Hammon2017) in preweaning calves.

Impact of colostrum supply on postnatal maturation

Colostrum supply and intestinal development and maturation

Bovine colostrum provides newborn calves with high amounts of nutrient and non-nutrient biologically active substances (Blum and Baumrucker, Reference Blum and Baumrucker2008; Nissen et al., Reference Nissen, Andersen, Bendixen, Ingvartsen and Rontved2017). In addition to the great importance of colostral immunoglobulins for the passive immunity of neonatal calves (Barrington and Parish, Reference Barrington and Parish2001; Godden, Reference Godden2008), colostrum contains a large number of immunomodulatory peptides that may also affect neonatal immune response (Chase et al., Reference Chase, Hurley and Reber2008; Stelwagen et al., Reference Stelwagen, Carpenter, Haigh, Hodgkinson and Wheeler2009; Nissen et al., Reference Nissen, Andersen, Bendixen, Ingvartsen and Rontved2017). Some of these factors are provided by colostral immune cells that are involved in the establishment of local and systemic neonatal immunity (Liebler-Tenorio et al., Reference Liebler-Tenorio, Riedel-Caspari and Pohlenz2002; Stelwagen et al., Reference Stelwagen, Carpenter, Haigh, Hodgkinson and Wheeler2009; Langel et al., Reference Langel, Wark, Garst, James, McGilliard, Petersson-Wolfe and Kanevsky-Mullarky2015). In addition, potential effects of colostrum on the neonatal microbiome in the gut are likely and become more important in the research of calf nutrition (Malmuthuge and Guan, Reference Malmuthuge and Guan2017). The importance of the colostrum supply for the development and maturation of the immune system of the newborn calf is far beyond the provision of immunoglobulins. Recent studies in humans, investigating the effects of breastmilk feeding on neonatal intestinal development, illustrate the significance of colostral immune cells and the intestinal microbiome on the maturation of the neonatal immune response in the gut (Molès et al., Reference Molès, Tuaillon, Kankasa, Bedin, Nagot, Marchant, McDermid and Van de Perre2018). A comparable function of bovine colostrum in the neonatal intestine of the calf is conceivable but requires more investigations.

Bovine colostrum has an overall importance for the postnatal development of the gut (Blum, Reference Blum2006). The high concentrations of hormones, growth factors and cell-modulating factors in colostrum (Blum and Baumrucker, Reference Blum and Baumrucker2008; Nissen et al., Reference Nissen, Andersen, Bendixen, Ingvartsen and Rontved2017) stimulate villus growth of the small intestinal mucosa in calves (Blum, Reference Blum2006; Steinhoff-Wagner et al., Reference Steinhoff-Wagner, Zitnan, Schönhusen, Pfannkuche, Hudakova, Metges and Hammon2014). Colostrum feeding promotes mucosal cell growth and protein synthesis in the enterocytes of neonatal mammals (Donovan and Odle, Reference Donovan and Odle1994; Burrin et al., Reference Burrin, Davis, Ebner, Schoknecht, Fiorotto, Reeds and McAvoy1995; Xu, Reference Xu1996). The amount of overall ingested colostrum corresponds to the villus size in the intestinal mucosa, leading to a greater villus size in repeatedly colostrum-fed calves (Blum, Reference Blum2006). When feeding a colostrum extract, that is, a fraction originating from first colostrum including most of the growth-promoting peptides, together with a milk-based formula, the villus size is stimulated when compared to a milk-based formula feeding with similar protein and energy as in colostrum but no growth-stimulating peptides (Roffler et al., Reference Roffler, Fäh, Sauter, Hammon, Gallmann, Brem and Blum2003). This finding supports the general assumption that colostral peptides, such as IGF-I, or hormones, such as insulin, are involved in the growth-stimulating effect on the intestinal mucosa of neonatal calves (Blum, Reference Blum2006).

In general, the proliferation rate of intestinal crypt cells depends on feeding (Johnson Reference Johnson1988; Mathers, Reference Mathers1998). Colostrum or colostral components stimulate crypt cell proliferation in the intestinal mucosa of neonatal calves (Blum, Reference Blum2006). When comparing colostrum feeding and milk-based formula feeding (same nutrient content but no growth-promoting bioactive factors as colostrum during the first 3 days after birth), the greater stimulation of cell proliferation corresponded to the greater villus growth in colostrum than formula-fed calves on day 8 of life (Blum, Reference Blum2006). The cell turnover of the intestinal mucosa depends on cell proliferation and programmed cell death (apoptosis; mainly seen at the villus tips) (Ramachandran et al., Reference Ramachandran, Madesh and Balasubramanian2000). Colostrum intake reduces apoptosis of epithelial cells and therefore prolongs the lifespan of the epithelial cells (Blum, Reference Blum2006).

Milk-borne factors such as IGF-I are known for their stimulation of cell proliferation (Burrin et al., Reference Burrin, Wester, Davis, Amick and Heath1996; Hammon et al., Reference Hammon, Steinhoff-Wagner, Flor, Schönhusen and Metges2013; Ontsouka et al., Reference Ontsouka, Albrecht and Bruckmaier2016) as well as inhibition of cell death due to apoptosis or inflammation in the intestinal mucosa (Mylonas et al., Reference Mylonas, Matsouka, Papandoniou, Vagianos, Kalfarentzos and Alexandrides2000; Blum, Reference Blum2006). Recombinant human IGF-I, fed together with MR in neonatal calves, increases intestinal cell proliferation (Blum, Reference Blum2006). A more distinct stimulation of mucosal cell proliferation is observed when feeding a colostrum extract (see above) instead of a single growth-promoting peptide (Roffler et al., Reference Roffler, Fäh, Sauter, Hammon, Gallmann, Brem and Blum2003). This finding indicates that not a single factor but the interaction of the large amount of growth-stimulating substances in the colostrum promotes intestinal cell proliferation and growth (Hammon et al., Reference Hammon, Steinhoff-Wagner, Schönhusen, Metges and Blum2012). Receptors for IGF-I, IGF-II and insulin (IGF1R, IGF2R and InsR, respectively) are present in the intestinal mucosa throughout the total gut in neonatal calves, and their expression and/or binding capacities are modified by colostrum feeding and orally administered rhIGF-I (Blum, Reference Blum2006; Hammon et al., Reference Hammon, Steinhoff-Wagner, Flor, Schönhusen and Metges2013; Ontsouka et al., Reference Ontsouka, Albrecht and Bruckmaier2016). The density of IGF1R and InsR, but not IGF2R, in the intestinal mucosa seems to be associated with crypt cell proliferation (Georgiev et al., Reference Georgiev, Georgieva, Pfaffl, Hammon and Blum2003).

Most biologically active factors in colostrum, such as IGF-I, IGF-II and insulin, are barely absorbed and therefore likely have no systemic function (Blum, Reference Blum2006; Hammon et al., Reference Hammon, Steinhoff-Wagner, Flor, Schönhusen and Metges2013). Therefore, local effects of colostral factors on crypt cell proliferation, intestinal epithelial growth and intestinal maturation may dominate in neonatal farm animals (Donovan and Odle, Reference Donovan and Odle1994; Reference XuXu, 1996; Blum, Reference Blum2006). However, recent findings in calves indicate the absorption of colostral adiponectin in neonatal calves (Kesser et al., Reference Kesser, Hill, Heinz, Koch, Rehage, Steinhoff-Wagner, Hammon, Mielenz, Sauerwein and Sadri2015), an adipokine involved in the regulation of insulin sensitivity (Kadowaki et al., Reference Kadowaki, Yamauchi, Kubota, Hara, Ueki and Tobe2006). With respect to the lactocrine signalling theory described in pigs (Bartol et al., Reference Bartol, Wiley, Miller, Silva, Roberts, Davolt, Chen, Frankshun, Camp, Rahman, Vallet and Bagnell2013), the intestinal absorption of adiponectin may contribute to the stimulation of anabolic metabolism in neonatal calves after colostrum feeding (Hammon et al., Reference Hammon, Steinhoff-Wagner, Schönhusen, Metges and Blum2012). Colostrum feeding supports protein synthesis in the skeletal muscle of piglets (Burrin et al., Reference Burrin, Davis, Ebner, Schoknecht, Fiorotto, Reeds and McAvoy1995), and the present research suggests a similar effect in neonatal calves, indicating enhanced protein synthesis in skeletal muscle due to insulin action after colostrum feeding (Hammon et al., Reference Hammon, Steinhoff-Wagner, Schönhusen, Metges and Blum2012; Sadri et al., Reference Sadri, Steinhoff-Wagner, Hammon, Bruckmaier, Gors and Sauerwein2017). The impact of adiponectin on this finding, however, is not clear and needs further investigation.

Colostrum supply and glucose metabolism

Due to its growth-stimulating effect in the small intestine, colostrum intake promotes the absorptive capacity of the small intestine. Measurements of xylose and glucose absorption in neonatal calves clearly indicate a greater absorption after feeding with colostrum instead of formula or MR (Blum, Reference Blum2006; Steinhoff-Wagner et al., Reference Steinhoff-Wagner, Gors, Junghans, Bruckmaier, Kanitz, Metges and Hammon2011; Gruse et al., Reference Gruse, Görs, Tuchscherer, Otten, Weitzel, Metges, Wolffram and Hammon2015). Xylose absorption on day 5 of age after feeding colostrum only once was similar to that after feeding colostrum for the first 3 days after birth (Hammon et al., Reference Hammon, Steinhoff-Wagner, Flor, Schönhusen and Metges2013). Therefore, the intake of first colostrum during the first hours after birth is of great importance for glucose absorption and the postnatal glucose status in neonatal calves. In contrast, digestive enzymes and mucosal transporters with respect to carbohydrate digestion, such as lactase and SGLT1 and GLUT2, seem to be less affected by colostrum feeding. A distinct stimulation of lactase activity and the glucose transporter when feeding colostrum instead of a milk-based formula was barely observed in neonatal calves (Sauter et al., Reference Sauter, Roffler, Philipona, Morel, Rome, Guilloteau, Blum and Hammon2004; Steinhoff-Wagner et al., Reference Steinhoff-Wagner, Zitnan, Schönhusen, Pfannkuche, Hudakova, Metges and Hammon2014). For more details concerning digestive enzymes in neonatal calves, readers are referred to Guilloteau et al. (Reference Guilloteau, Zabielski and Blum2009a). Recent studies using metabolomics approaches in neonatal calves indicate that the uptake and metabolism of other nutrients (e.g. amino acids) are also influenced by colostrum feeding (Qi et al., Reference Qi, Zhao, Huang, Pan, Yang, Zhao, Hu and Cheng2018; Zhao et al., Reference Zhao, Qi, Huang, Pan, Cheng, Zhao and Yang2018).

First-pass glucose uptake in the splanchnic tissue on days 2 and 7 of life is greater in formula-fed than colostrum-fed calves, indicating a greater glucose utilisation in the splanchnic tissue (gastrointestinal tract and liver) of calves not fed with colostrum (Steinhoff-Wagner et al., Reference Steinhoff-Wagner, Gors, Junghans, Bruckmaier, Kanitz, Metges and Hammon2011; Gruse et al., Reference Gruse, Görs, Tuchscherer, Otten, Weitzel, Metges, Wolffram and Hammon2015). Possibly, nutrient absorption is generally impaired in formula-fed calves, leading to increased glucose utilisation in the splanchnic tissue, whereas colostrum-fed calves are able to use greater amounts of digested fat and protein as energy fuel in the splanchnic tissue. This hypothesis is supported by the finding that oral fat absorption is greater in colostrum- than in formula-fed or MR-fed calves, providing more fat (i.e. medium-chain fatty acids from colostrum) as energy fuel in the splanchnic tissue that partly can replace glucose utilisation (Hammon et al., Reference Hammon, Steinhoff-Wagner, Schönhusen, Metges and Blum2012). In contrast to intestinal glucose absorption, a stimulating influence of colostrum intake on endogenous glucose production, as supposed to be the case in piglets (Lepine et al., Reference Lepine, Boyd and Whitehead1991), does not occur in bovine neonates. Thus, growth-promoting substances of ingested colostrum do not affect endogenous glucose production in neonatal calves (Steinhoff-Wagner et al., Reference Steinhoff-Wagner, Gors, Junghans, Bruckmaier, Kanitz, Metges and Hammon2011). Nevertheless, the increased plasma glucose concentration and the greater hepatic glycogen content in colostrum-fed calves indicate an improved glucose status by colostrum feeding (Steinhoff-Wagner et al., Reference Steinhoff-Wagner, Gors, Junghans, Bruckmaier, Kanitz, Metges and Hammon2011; Hammon et al., Reference Hammon, Steinhoff-Wagner, Schönhusen, Metges and Blum2012 and Reference Hammon, Steinhoff-Wagner, Flor, Schönhusen and Metges2013). These findings lead to the conclusion that the improved glucose status in calves fed colostrum immediately after birth and for 3 days is a result of enhanced glucose absorption and probably of less glucose utilisation in the splanchnic tissue but is not a result of increased endogenous glucose production.

Colostrum supply and maturation in the somatotropic axis

The elevated glucose availability and the improved insulin status in colostrum-fed calves are important prerequisites for the accelerated maturation of the somatotropic axis, as indicated by several studies in neonatal calves that were previously summarised (Blum, Reference Blum2006; Hammon et al., Reference Hammon, Steinhoff-Wagner, Schönhusen, Metges and Blum2012). The stimulation of gastrointestinal hormones due to colostrum feeding may contribute to the elevated insulin secretion in the calves (Hadorn et al., Reference Hadorn, Hammon, Bruckmaier and Blum1997; Inabu et al., Reference Inabu, Pyo, Pletts, Guan, Steele and Sugino2019). The elevated insulin status due to colostrum feeding in neonatal calves is probably the trigger for stimulating endogenous IGF-I and the postnatal somatotropic axis (Hammon et al., Reference Hammon, Steinhoff-Wagner, Schönhusen, Metges and Blum2012) because glucose and insulin stimulate the hepatic gene expression of the growth hormone receptor and IGF-I as well as IGF-I secretion (Brameld et al., Reference Brameld, Gilmour and Buttery1999; Butler et al., Reference Butler, Marr, Pelton, Radcliff, Lucy and Butler2003). On the other hand and as discussed earlier, studies in neonatal calves and piglets indicate no intestinal absorption of colostral IGF-I or insulin (Donovan et al., Reference Donovan, Chao, Zijlstra and Odle1997; Blum, Reference Blum2006). Thus, the endogenously produced IGF-I determines the IGF-I status of the calf. Therefore, the nutrient supply is responsible for the maturation of the neonatal somatotropic axis, and the colostral IGF-I promotes intestinal development of the neonatal calf but does not contribute to systemic IGF-I availability (Blum, Reference Blum2006; Hammon et al., Reference Hammon, Steinhoff-Wagner, Schönhusen, Metges and Blum2012; Ontsouka et al., Reference Ontsouka, Albrecht and Bruckmaier2016).

In summary, the postnatal maturation of the neonatal intestine is enhanced due to colostrum intake, and the improved intestinal maturation results in a greater nutrient absorption and stimulation of anabolic processes that are a prerequisite for accelerated postnatal growth.

Development of the preweaning calf due to intensive milk feeding

Definition of intensive milk feeding

After the colostrum period, the calf depends on the intake of liquid feed in the form of milk or high-quality MR for nutrient supply. Although it is a common feeding strategy to increase solid feed intake as soon as possible in the preweaning period by reducing milk feeding (Huber, Reference Huber1969; Gelsinger et al., Reference Gelsinger, Heinrichs and Jones2016; Kertz et al., Reference Kertz, Hill, Quigley, Heinrichs, Linn and Drackley2017), solid feed intake during the first 3 weeks of age is low, and the digestion of solid feed is impaired due to the immature forestomach in the postnatal period (Drackley, Reference Drackley2008; Khan et al., Reference Khan, Bach, Weary and von Keyserlingk2016; Frieten et al., Reference Frieten, Gerbert, Koch, Dusel, Eder, Kanitz, Weitzel and Hammon2017). Thus, sufficient milk or MR supply during the first weeks of life is a prerequisite for calf growth and development. The World Organisation for Animal Health (OIE) defines animal welfare in the Terrestrial Animal Health Code as a state where the animal is healthy, comfortable, well nourished, safe, able to express natural behaviour and not suffering from pain, fear and distress (OIE, 2017). Feeding calves limited amounts of liquid feed (i.e. 4 to 6 kg/day) during the first weeks of life results in a lack of expression in natural suckling behaviour (Schiessler et al., Reference Schiessler, Nussbaum, Hammon and Blum2002; Miller-Cushon and DeVries, Reference Miller-Cushon and DeVries2015) followed by hunger (Jensen and Holm, Reference Jensen and Holm2003; De Paula Vieira et al., Reference De Paula Vieira, Guesdon, De Passille, von Keyserlingk and Weary2008; Borderas et al., Reference Borderas, de Passille and Rushen2009; Gerbert et al., Reference Gerbert, Frieten, Koch, Dusel, Eder, Stefaniak, Bajzert, Jawor, Tuchscherer and Hammon2018) and stress for the calves. Allowing calves to drink unlimited amounts of milk or MR for several weeks during the preweaning period more than doubles liquid feed intake compared with restricted amounts of 4 to 6 kg/day of MR or milk (Hammon et al., Reference Hammon, Schiessler, Nussbaum and Blum2002; Maccari et al., Reference Maccari, Wiedemann, Kunz, Piechotta, Sanftleben and Kaske2015; Schäff et al., Reference Schäff, Gruse, Maciej, Mielenz, Wirthgen, Hoeflich, Schmicke, Pfuhl, Jawor, Stefaniak and Hammon2016; Frieten et al., Reference Frieten, Gerbert, Koch, Dusel, Eder, Kanitz, Weitzel and Hammon2017). Therefore, an intensive milk-feeding programme contributes to the overall well-being of preweaning calves (Von Keyserlingk et al., Reference Von Keyserlingk, Rushen, de Passille and Weary2009; FAWC, 2015; OIE, 2017).

An intensive milk feeding provides milk or MR in unrestricted amounts all day long for 24 h. The calves have ad libitum access to milk (Jasper and Weary, Reference Jasper and Weary2002; Schiessler et al., Reference Schiessler, Nussbaum, Hammon and Blum2002; Maccari et al., Reference Maccari, Wiedemann, Kunz, Piechotta, Sanftleben and Kaske2015) or MR feeding (Schäff et al., Reference Schäff, Gruse, Maciej, Mielenz, Wirthgen, Hoeflich, Schmicke, Pfuhl, Jawor, Stefaniak and Hammon2016; Frieten et al., Reference Frieten, Gerbert, Koch, Dusel, Eder, Kanitz, Weitzel and Hammon2017; Korst et al., Reference Korst, Koch, Kesser, Müller, Romberg, Rehage, Eder and Sauerwein2017) for several weeks. Ad libitum MR feeding (125 g powder/l, 21.7% CP and 18.3 MJ metabolisable energy (ME) per kg DM) provides on average 1.6 kg DM/day, 35 MJ ME/day and 347 g protein/day to the calves during the intensive milk-feeding period. In this context, daily peaks of more than 2 kg DM intake were observed in previous studies (Schäff et al., Reference Schäff, Gruse, Maciej, Mielenz, Wirthgen, Hoeflich, Schmicke, Pfuhl, Jawor, Stefaniak and Hammon2016; Frieten et al., Reference Frieten, Gerbert, Koch, Dusel, Eder, Kanitz, Weitzel and Hammon2017). Such feeding schedules provide much more protein and energy than commonly used milk-feeding programmes of 4 to 6 kg milk/day. Ad libitum milk-feeding programmes are comparable to early calf rearing in beef production where calves live together with their dams and have free access to milk all day long (Egli and Blum, Reference Egli and Blum1998; Schiessler et al., Reference Schiessler, Nussbaum, Hammon and Blum2002). Main findings of an intensive milk feeding protocol compared to restricted milk feeding on calf growth, organ development, metabolic and endocrine changes, feeding behaviour and immune response have been summarised in Table 1.

Table 1 Effects of intensive milk feeding immediately after birth on growth, development, behaviour and immune response in preweaning calves1

1 Intensive milk feeding is defined as daily milk or milk replacer intake of 20% of BW, ad libitum milk or milk replacer feeding or feeding enhanced amounts of milk replacer with elevated CP and fat content.

2 When not stated in the table, milk replacer contained 21% to 23% of CP and 17% to 20% of crude fat based on DM. Whole milk contained 320 to 350 g CP and 370 to 400 g crude fat/kg milk.

3 Data apply only for Holstein calves.

4 Two studies in the reference.

Stimulation of growth and endocrine growth regulation by intensive milk feeding

Ad libitum milk or MR feeding (Maccari et al., Reference Maccari, Wiedemann, Kunz, Piechotta, Sanftleben and Kaske2015; Schäff et al., Reference Schäff, Gruse, Maciej, Mielenz, Wirthgen, Hoeflich, Schmicke, Pfuhl, Jawor, Stefaniak and Hammon2016; Korst et al., Reference Korst, Koch, Kesser, Müller, Romberg, Rehage, Eder and Sauerwein2017) or enhanced milk-feeding programmes using MR with a greater protein content (up to 30% CP in DM) (Smith et al., Reference Smith, Van Amburgh, Diaz, Lucy and Bauman2002; Geiger et al., Reference Geiger, Parsons, James and Akers2016) resulted in an elevated body growth during the preweaning period when compared to restricted milk or MR feeding (4 to 6 kg milk or MR/day). In addition to stimulating muscle and fat growth (Bartlett et al., Reference Bartlett, McKeith, VandeHaar, Dahl and Drackley2006; Geiger et al., Reference Geiger, Parsons, James and Akers2016; Schäff et al., Reference Schäff, Gruse, Maciej, Mielenz, Wirthgen, Hoeflich, Schmicke, Pfuhl, Jawor, Stefaniak and Hammon2016; Koch et al., Reference Koch, Gerbert, Frieten, Dusel, Eder, Zitnan and Hammon2019), intensive milk or MR intake accelerates organ growth, for example, small intestine, mammary gland, thymus and endocrine pancreas (Prokop et al., Reference Prokop, Kaske, Maccari, Lucius, Kunz and Wiedemann2015; Geiger et al., Reference Geiger, Parsons, James and Akers2016; Soberon and Van Amburgh, Reference Soberon and Van Amburgh2017; Koch et al., Reference Koch, Gerbert, Frieten, Dusel, Eder, Zitnan and Hammon2019). As discussed later in this review, the velocity of body growth could temporarily decrease in intensive milk-fed calves during the weaning process due to the adaptation to solid feed intake. However, BW at the end of the weaning process is still greater in intensively than restrictively milk-fed calves (Maccari et al., Reference Maccari, Wiedemann, Kunz, Piechotta, Sanftleben and Kaske2015; Schäff et al., Reference Schäff, Gruse, Maciej, Mielenz, Wirthgen, Hoeflich, Schmicke, Pfuhl, Jawor, Stefaniak and Hammon2016; Frieten et al., Reference Frieten, Gerbert, Koch, Dusel, Eder, Kanitz, Weitzel and Hammon2017).

Studies on the hepatic transcriptome and proteome in lambs and metabolome in blood plasma of calves reveal marked changes with respect to protein and energy metabolism when animals receive MR ad libitum instead of in restricted amounts (Kenéz et al., Reference Kenéz, Koch, Korst, Kesser, Eder, Sauerwein and Huber2018; Santos et al., Reference Santos, Giraldez, Frutos and Andres2019). In lambs, restricted MR feeding stimulates hepatic pathways involved in gluconeogenesis, amino acid degradation and hepatic fatty acid oxidation, which points at changes in energy utilisation to stabilise glucose homeostasis as compared to ad libitum MR-fed lambs (Santos et al., Reference Santos, Giraldez, Frutos and Andres2019). In calves, ad libitum instead of restricted MR feeding seems to increase the capacity for mitochondrial transport of fatty acids and probably affects fatty acid oxidation as well (Kenéz et al., Reference Kenéz, Koch, Korst, Kesser, Eder, Sauerwein and Huber2018). In addition, an enhanced MR-feeding programme leads to a greater metabolic activity in muscle and fat tissue as well as the ruminal epithelium (Naeem et al., Reference Naeem, Drackley, Stamey and Loor2012 and Reference Naeem, Drackley, Lanier, Everts, Rodriguez-Zas and Loor2014; Wang et al., Reference Wang, Drackley, Stamey-Lanier, Keisler and Loor2014; Leal et al., Reference Leal, Romao, Hooiveld, Soberon, Berends, Boekshoten, Van Amburgh, Martin-Tereso and Steele2018).

The improved growth development and protein accretion in calves fed intensively with milk or MR are confirmed by the stimulation of the somatotropic axis (Maccari et al., Reference Maccari, Wiedemann, Kunz, Piechotta, Sanftleben and Kaske2015; Schäff et al., Reference Schäff, Gruse, Maciej, Mielenz, Wirthgen, Hoeflich, Schmicke, Pfuhl, Jawor, Stefaniak and Hammon2016; Frieten et al., Reference Frieten, Gerbert, Koch, Dusel, Eder, Hoeflich, Mielenz and Hammon2018; Haisan et al., Reference Haisan, Oba, Ambrose and Steele2018). Important elements of the somatotropic axis are growth hormone (GH), IGF-I and several IGF-binding proteins (IGFBPs). The postnatal interaction of GH, IGF-I and IGFBP affects body growth and organ development in mammals (Breier et al., Reference Breier, Oliver, Gallaher and Cronjé2000), including the development of the mammary gland (Akers, Reference Akers2006) and immune function (Clark, Reference Clark1997). The stimulation of the postnatal somatotropic axis depends on the nutrient supply and reflects the glucose and insulin status of the animal (Brameld et al., Reference Brameld, Gilmour and Buttery1999; Renaville et al., Reference Renaville, Van Eenaeme, Breier, Vleurick, Bertozzi, Gengler, Hornick, Parmentier, Istasse, Haezebroeck, Massart and Portetelle2000; Smith et al., Reference Smith, Van Amburgh, Diaz, Lucy and Bauman2002). Plasma IGF-I and IGFBP-3 concentrations are elevated, and the IGFBP-2 concentration is decreased during growth in well-nourished animals as compared to animals of same age with restricted feed intake (Breier et al., Reference Breier, Oliver, Gallaher and Cronjé2000; Renaville et al., Reference Renaville, Van Eenaeme, Breier, Vleurick, Bertozzi, Gengler, Hornick, Parmentier, Istasse, Haezebroeck, Massart and Portetelle2000). A key factor in maturation of the somatotropic axis is the increased expression of the GH receptor, particularly in the liver, with age (Breier et al., Reference Breier, Oliver, Gallaher and Cronjé2000; Hammon et al., Reference Hammon, Steinhoff-Wagner, Schönhusen, Metges and Blum2012). The GH receptor mediates GH action on IGF-I synthesis and secretion and is stimulated by insulin (Breier et al., Reference Breier, Oliver, Gallaher and Cronjé2000; Butler et al., Reference Butler, Marr, Pelton, Radcliff, Lucy and Butler2003). The glucose, insulin, IGF-I and IGFBP-3 plasma concentrations are much greater, and hepatic gene expression of the GH receptor and IGF-I is higher in intensively milk-fed calves than in calves with restricted milk intake (e.g., 6 l MR/day; Maccari et al., Reference Maccari, Wiedemann, Kunz, Piechotta, Sanftleben and Kaske2015; Schäff et al., Reference Schäff, Gruse, Maciej, Mielenz, Wirthgen, Hoeflich, Schmicke, Pfuhl, Jawor, Stefaniak and Hammon2016; Frieten et al., Reference Frieten, Gerbert, Koch, Dusel, Eder, Kanitz, Weitzel and Hammon2017 and Reference Frieten, Gerbert, Koch, Dusel, Eder, Hoeflich, Mielenz and Hammon2018). The IGFBP-2 plasma concentration and hepatic gene expression behave the other way round, as expected from the literature (Renaville et al., Reference Renaville, Van Eenaeme, Breier, Vleurick, Bertozzi, Gengler, Hornick, Parmentier, Istasse, Haezebroeck, Massart and Portetelle2000; Schäff et al., Reference Schäff, Gruse, Maciej, Mielenz, Wirthgen, Hoeflich, Schmicke, Pfuhl, Jawor, Stefaniak and Hammon2016; Frieten et al., Reference Frieten, Gerbert, Koch, Dusel, Eder, Hoeflich, Mielenz and Hammon2018). No signs of impaired insulin response are seen during enhanced milk feeding in calves (MacPherson et al., Reference MacPherson, Meale, Macmillan, Haisan, Bench, Oba and Steele2019).

During the first weeks of life, the elevated concentrate intake in restrictively milk-fed calves cannot compensate for impaired nutrient intake due to reduced milk feeding, and consequently, the somatotropic axis is not stimulated during early postnatal life when concentrate and forage feeding are favoured instead of milk feeding. In particular, the elevated IGFBP-2 plasma concentration in milk-restricted-fed calves indicates an impaired nutrient intake (Renaville et al., Reference Renaville, Van Eenaeme, Breier, Vleurick, Bertozzi, Gengler, Hornick, Parmentier, Istasse, Haezebroeck, Massart and Portetelle2000; Schäff et al., Reference Schäff, Gruse, Maciej, Mielenz, Wirthgen, Hoeflich, Schmicke, Pfuhl, Jawor, Stefaniak and Hammon2016; Frieten et al., Reference Frieten, Gerbert, Koch, Dusel, Eder, Hoeflich, Mielenz and Hammon2018). An impaired nutrient supply with decreased IGF-I and IGFBP-3 and increased IGFBP-2 plasma concentrations also occurs during weaning when milk feeding is reduced too quickly and the solid feed intake does not meet nutrition requirements (Schäff et al., Reference Schäff, Gruse, Maciej, Mielenz, Wirthgen, Hoeflich, Schmicke, Pfuhl, Jawor, Stefaniak and Hammon2016; Frieten et al., Reference Frieten, Gerbert, Koch, Dusel, Eder, Hoeflich, Mielenz and Hammon2018). These changes in the somatotropic axis are reflected by a depressed growth rate during the weaning process (Schäff et al., Reference Schäff, Gruse, Maciej, Mielenz, Wirthgen, Hoeflich, Schmicke, Pfuhl, Jawor, Stefaniak and Hammon2016; Frieten et al., Reference Frieten, Gerbert, Koch, Dusel, Eder, Kanitz, Weitzel and Hammon2017). To prevent a growth depression during weaning in calves with an intensified milk-feeding programme, a delayed weaning age or individual weaning based on solid feed intake is recommended (Eckert et al., Reference Eckert, Brown, Leslie, DeVries and Steele2015; de Passillé and Rushen, Reference de Passillé and Rushen2016; Welboren et al., Reference Welboren, Leal, Steele, Khan and Martin-Tereso2019). Parameters of the somatotropic axis may provide useful information on the metabolic status and may help to avoid detrimental weaning programmes in calves.

Development and maturation of the gastrointestinal tract and immune response by intensive milk feeding

An intensive milk feeding strategy affects intestinal development. The size and the absorptive capacity increase in preweaning calves with an enhanced MR-feeding programme (Geiger et al., Reference Geiger, Parsons, James and Akers2016; Koch et al., Reference Koch, Gerbert, Frieten, Dusel, Eder, Zitnan and Hammon2019). In addition, an intensive milk or MR-feeding programme seems to stimulate the expression of long non-coding RNA involved in the regulation of tight-junction protein synthesis (Weikard et al., Reference Weikard, Hadlich, Hammon, Frieten, Gerbert, Koch, Dusel and Kuehn2018). It is well established in ruminants that the diet affects tight-junction protein expression (Steele et al., Reference Steele, Penner, Chaucheyras-Durand and Guan2016). According to the upregulation of tight-junction protein-encoding genes, it is suggested that the first week of life is crucial for the development of the intestinal epithelium and the intestinal barrier of the mucosal immune system in calves (Malmuthuge and Guan, Reference Malmuthuge and Guan2017). In piglets, specific amino acids can affect intestinal permeability and integrity, protein synthesis, intestinal repair after injury and cell proliferation in the gastrointestinal tract (Jacobi and Odle, Reference Jacobi and Odle2012). Because of its composition and ingredients, milk provide the best conditions for nutrient supply in the postnatal and preweaning period to promote intestinal integrity in the bovine (Steele et al., Reference Steele, Penner, Chaucheyras-Durand and Guan2016).

There is growing evidence that an adequate nutrient supply is important for the maturation of the intestinal immune system and for successful defence against pathogens (Khan et al., Reference Khan, Weary and von Keyserlingk2011; Hammon et al., Reference Hammon, Frieten, Gerbert, Koch, Dusel, Weikard and Kühn2018). A greater nutrient supply may have beneficial effects on intestinal maturation, including the generation of a proper adaptive immune system and a stable microbiota, which may protect against diarrhoeal diseases in the neonatal and preweaning period. Common feeding schedules of not more than 6 kg of milk per day may delay the establishment of a proper immune response and microbiota in the intestine. Studies in neonatal calves have investigated the diet-dependent establishment of the intestinal microbiome, but the impact of the milk amount on the intestinal microbiome is still unclear (Malmuthuge and Guan, Reference Malmuthuge and Guan2017). A higher plane (considering the amount of MR, concentration of MR and protein and fat concentration) of nutrition seems to protect the intestine against pathogenic infections and promote overcoming of pathogenic infections. Calves with a higher plane of nutrition (intake energy: 1.3 MJ/kg metabolic BW instead of 0.5 MJ/kg metabolic BW by MR feeding) indicate a faster resolution from diarrhoea caused by infection with Cryptosporidium parvum (Ollivett et al., Reference Ollivett, Nydam, Linden, Bowman and Van Amburgh2012). Ballou et al. (Reference Ballou, Hanson, Cobb, Obeidat, Sellers, Pepper-Yowell, Carroll, Earleywine and Lawhon2015) showed that a higher plane of nutrition (610 and 735 g/day DM MR during week 1 and weeks 2 to 6, respectively, of a 28% CP and 25% fat MR instead of 409 g/day DM MR of a 20% CP and 20% fat MR) results in a higher resistance against Salmonella typhimurium in postweaning calves. Even there is first evidence that an intensive milk or MR feeding stimulates intestinal development and maturation and the intestinal immune response (Reference Hammon, Frieten, Gerbert, Koch, Dusel, Weikard and KühnHammon et al., 2018), more studies are needed to investigate the interaction between the changes of the intestinal microbiome and immune response due to an intensive milk-feeding programme.

An intensive milk feeding may delay rumen development by reducing solid feed intake during the early preweaning period (Baldwin et al., Reference Baldwin, McLeod, Klotz and Heitmann2004; Khan et al., Reference Khan, Weary and von Keyserlingk2011 and Reference Khan, Bach, Weary and von Keyserlingk2016). However, solid feed intake and rumen development accelerate during the weaning process, and rumen function is not impaired at the end of the weaning process when calves received 20% instead of 10% of BW milk per day (Khan et al., Reference Khan, Weary and von Keyserlingk2011). Rumen papilla growth and concentrations of volatile fatty acids are the same when calves are fed MR ad libitum for 5 and 8 weeks after birth, respectively, compared to 6 kg/day MR intake (Schäff et al., Reference Schäff, Gruse, Maciej, Pfuhl, Zitnan, Rajsky and Hammon2018; Koch et al., Reference Koch, Gerbert, Frieten, Dusel, Eder, Zitnan and Hammon2019). These findings are supported by the fact that ad libitum milk-fed calves immediately increase their solid feed intake and plasma β-hydroxybutyrate concentration in blood when MR intake is reduced (Schäff et al., Reference Schäff, Gruse, Maciej, Mielenz, Wirthgen, Hoeflich, Schmicke, Pfuhl, Jawor, Stefaniak and Hammon2016; Frieten et al., Reference Frieten, Gerbert, Koch, Dusel, Eder, Kanitz, Weitzel and Hammon2017; Welboren et al., Reference Welboren, Leal, Steele, Khan and Martin-Tereso2019). Plasma β-hydroxybutyrate results from ketogenesis from butyrate in the rumen epithelial cells and is an indicator for maturation of the rumen function (Baldwin et al., Reference Baldwin, McLeod, Klotz and Heitmann2004). Interestingly, the metabolic activity in ruminal epithelium is enhanced in calves fed elevated amounts of MR (Naeem et al., Reference Naeem, Drackley, Stamey and Loor2012 and Reference Naeem, Drackley, Lanier, Everts, Rodriguez-Zas and Loor2014).

In summary, an intensive milk-feeding regime is required to realise the potential for growth and development in preweaning calves. Body growth and organ maturation are improved with an intensive milk-feeding programme, and calves are less hungry and probably more resilient to infectious diseases during the preweaning period. The development of the forestomach could be delayed during the intensive milk-feeding period, but applying an adapted weaning protocol for intensive milk-fed calves avoids an impaired rumen development and body growth depression during the postweaning period.

Impact of preweaning growth and development on lifetime performance and health

Colostrum and milk feeding not only influence the postnatal and preweaning development and growth of the calves but also influence performance and health in later life (Van Amburgh und Soberon, Reference Van Amburgh and Soberon2013; Huber, Reference Huber2018). The improved mammary gland development during the preweaning rearing period is an example of the importance of the nutrient supply during the preweaning period for organ development (Geiger et al., Reference Geiger, Parsons, James and Akers2016; Soberon and Van Amburgh, Reference Soberon and Van Amburgh2017), with consequences for lifetime performance (Van Amburgh und Soberon, Reference Van Amburgh and Soberon2013). However, presently it is not known whether early calf nutrition has long-lasting effects on other organ systems, for example, the liver or the immune system. Culling rates are still high in dairy production, and metabolic diseases and immune suppression around calving are heavily involved in this unfavourable situation (Hare et al., Reference Hare, Norman and Wright2006; Ingvartsen and Moyes, Reference Ingvartsen and Moyes2015; Probo et al., Reference Probo, Pascottini, LeBlanc, Opsomer and Hostens2018; Gross and Bruckmaier, Reference Gross and Bruckmaier2019). Interestingly, different patterns of metabolic parameters due to variable milk or MR feeding in preweaning calves seem to be maintained when calves become dairy cows, and epigenetic effects due to different milk-feeding programmes during the preweaning period have been assumed (Kenéz et al., Reference Kenéz, Koch, Korst, Kesser, Eder, Sauerwein and Huber2018). Postnatal nutritional programming is well known from human studies and in other species (Guilloteau et al., Reference Guilloteau, Zabielski, Hammon and Metges2009b), but it still remains unclear whether variable metabolic profiles of young individuals are conserved and are the basis for the different metabolic types of the adults in their productive life span. Thus, more research is needed to investigate the impact of early calf nutrition on metabolic performance in later life and whether early calf nutrition may improve robustness and resilience in dairy cows.

Conclusions

An intensive milk-feeding programme, starting immediately after birth, with an enhanced colostrum intake and subsequent intensive milk feeding supports postnatal growth and development of dairy calves, prevents behavioural anomalies and promotes the raising of robust young animals. Providing only 4 to 6 kg milk/day to the preweaning calves is not consistent with animal welfare principles (FAWC, 2015; OIE, 2017; Huber, Reference Huber2018). Thus, a change of the early calf management is needed to follow the natural processes of preweaning calf rearing, for example, as known from beef calf management (Egli and Blum, Reference Egli and Blum1998; Schiessler et al., Reference Schiessler, Nussbaum, Hammon and Blum2002). Research will continue to investigate the impact of an intensive milk-feeding regime on raising robust and well-performing dairy cows and bulls.

Acknowledgements

This review is based on an invited presentation at the 13th International Symposium on Ruminant Physiology (ISRP 2019) held in Leipzig, Germany, September 2019.

H. M. Hammon 0000-0001-8698-1257

Declaration of interest

The authors declare no conflicts of interest.

Ethics statement

None.

Software and data repository resources

None.

References

Akers, RM 2006. Major advances associated with hormone and growth factor regulation of mammary growth and lactation in dairy cows. Journal of Dairy Science 89, 12221234.CrossRefGoogle ScholarPubMed
Baldwin, RL, McLeod, KR, Klotz, JL and Heitmann, RN 2004. Rumen development, intestinal growth and hepatic metabolism in the pre- and postweaning ruminant. Journal of Dairy Science 87, E55E65.CrossRefGoogle Scholar
Ballou, MA 2012. Immune responses of Holstein and Jersey calves during the preweaning and immediate postweaned periods when fed varying planes of milk replacer. Journal of Dairy Science 95, 73197330.CrossRefGoogle ScholarPubMed
Ballou, MA, Hanson, DL, Cobb, CJ, Obeidat, BS, Sellers, MD, Pepper-Yowell, AR, Carroll, JA, Earleywine, TJ and Lawhon, SD 2015. Plane of nutrition influences the performance, innate leukocyte responses, and resistance to an oral Salmonella enterica serotype Typhimurium challenge in Jersey calves. Journal of Dairy Science 98, 19721982.CrossRefGoogle Scholar
Barrington, GM and Parish, SM 2001. Bovine neonatal immunology. Veterinary Clinics of North America: Food Animal Practice 17, 463476.Google ScholarPubMed
Bartlett, KS, McKeith, FK, VandeHaar, MJ, Dahl, GE and Drackley, JK 2006. Growth and body composition of dairy calves fed milk replacers containing different amounts of protein at two feeding rates. Journal of Animal Science 84, 14541467.CrossRefGoogle ScholarPubMed
Bartol, FF, Wiley, AA, Miller, DJ, Silva, AJ, Roberts, KE, Davolt, ML, Chen, JC, Frankshun, AL, Camp, ME, Rahman, KM, Vallet, JL and Bagnell, CA 2013. Lactation biology symposium: lactocrine signaling and developmental programming. Journal of Animal Science 91, 696705.CrossRefGoogle ScholarPubMed
Blum, JW 2006. Nutritional physiology of neonatal calves. Journal of Animal Physiology and Animal Nutrition 90, 111.CrossRefGoogle ScholarPubMed
Blum, JW and Baumrucker, CR 2008. Insulin-like growth factors (IGFs), IGF binding proteins, and other endocrine factors in milk: role in the newborn. Advances in Experimental Medicine and Biology 606, 397422.CrossRefGoogle ScholarPubMed
Borderas, TF, de Passille, AMB and Rushen, J 2009. Feeding behavior of calves fed small or large amounts of milk. Journal of Dairy Science 92, 28432852.CrossRefGoogle ScholarPubMed
Brameld, JM, Gilmour, RS and Buttery, PJ 1999. Glucose and amino acids interact with hormones to control expression of insulin-like growth factor-I and growth hormone receptor mRNA in cultured pig hepatocytes. Journal of Nutrition 129, 12981306.CrossRefGoogle ScholarPubMed
Breier, BH, Oliver, MH and Gallaher, BW 2000. Regulation of growth and metabolism during postnatal development. In Ruminant physiology: digestion, metabolism, growth and reproduction (ed. Cronjé, PB 2000. Regulation of growth and metabolism during postnatal development. In Ruminant physiology: digestion, metabolism, growth and reproduction (ed. ), pp. 187204. CABI Publishing, New York, NY, USA.CrossRefGoogle Scholar
Burrin, DG, Davis, TA, Ebner, S, Schoknecht, PA, Fiorotto, ML, Reeds, PJ and McAvoy, S 1995. Nutrient-independent and nutrient-dependent factors stimulate protein synthesis in colostrum-fed newborn pigs. Pediatric Research 37, 593599.CrossRefGoogle ScholarPubMed
Burrin, DG, Wester, TJ, Davis, TA, Amick, S and Heath, JP 1996. Orally administered IGF-I increases intestinal mucosal growth in formula-fed neonatal pigs. American Journal of Physiology 270, R10851091.Google ScholarPubMed
Butler, ST, Marr, AL, Pelton, SH, Radcliff, RP, Lucy, MC and Butler, WR 2003. Insulin restores GH responsiveness during lactation-induced negative energy balance in dairy cattle: effects on expression of IGF-I and GH receptor 1A. Journal of Endocrinology 176, 205217.CrossRefGoogle ScholarPubMed
Chase, CCL, Hurley, DJ and Reber, AJ 2008. Neonatal immune development in the calf and its impact on vaccine response. Veterinary Clinics of North America: Food Animal Practice 24, 87104.Google ScholarPubMed
Clark, R 1997. The somatogenic hormones and insulin-like growth factor-1: stimulators of lymphopoiesis and immune function. Endocrine Reviews 18, 157179.CrossRefGoogle ScholarPubMed
Compton, CWR, Heuer, C, Thomsen, PT, Carpenter, TE, Phyn, CVC and McDougall, S 2017. Invited review: a systematic literature review and meta-analysis of mortality and culling in dairy cattle. Journal of Dairy Science 100, 116.CrossRefGoogle ScholarPubMed
Daniels, KM, Capuco, AV, McGilliard, ML, James, RE and Akers, RM 2009. Effects of milk replacer formulation on measures of mammary growth and composition in Holstein heifers. Journal of Dairy Science 92, 59375950.CrossRefGoogle ScholarPubMed
Daniels, KM, Hill, SR, Knowlton, KF, James, RE, McGilliard, ML and Akers, RM 2008. Effects of milk replacer composition on selected blood metabolites and hormones in preweaned Holstein heifers. Journal of Dairy Science 91, 26282640.CrossRefGoogle ScholarPubMed
Davis Rincker, LE, Vandehaar, MJ, Wolf, CA, Liesman, JS, Chapin, LT and Weber Nielsen, MS 2011. Effect of intensified feeding of heifer calves on growth, pubertal age, calving age, milk yield, and economics. Journal of Dairy Science 94, 35543567.CrossRefGoogle ScholarPubMed
de Passillé, AM and Rushen, J 2016. Using automated feeders to wean calves fed large amounts of milk according to their ability to eat solid feed. Journal of Dairy Science 99, 35783583.CrossRefGoogle ScholarPubMed
Department for Environment, Food and Rural Affairs (DEFRA) 2003. Improving calf survival. Retrieved on 20 June 2019 from http://adlib.everysite.co.uk/resources/000/020/709/calfsurvival.pdfGoogle Scholar
De Paula Vieira, A, Guesdon, V, De Passille, AM, von Keyserlingk, MAG and Weary, DM 2008. Behavioural indicators of hunger in dairy calves. Applied Animal Behaviour Science 109, 180189.CrossRefGoogle Scholar
Donovan, SM, Chao, JCJ, Zijlstra, RT and Odle, J 1997. Orally administered iodinated recombinant human insulin-like growth factor-I (I-125-rhIGF-I) is poorly absorbed by the newborn piglet. Journal of Pediatric Gastroenterology and Nutrition 24, 174182.CrossRefGoogle Scholar
Donovan, SM and Odle, J 1994. Growth factors in milk as mediators of infant development. Annual Review of Nutrition 14, 147167.CrossRefGoogle ScholarPubMed
Drackley, JK 2008. Calf nutrition from birth to breeding. Veterinary Clinics of North America: Food Animal Practice 24, 5586.Google ScholarPubMed
Eckert, E, Brown, HE, Leslie, KE, DeVries, TJ and Steele, MA 2015. Weaning age affects growth, feed intake, gastrointestinal development, and behavior in Holstein calves fed an elevated plane of nutrition during the preweaning stage. Journal of Dairy Science 98, 63156326.CrossRefGoogle ScholarPubMed
Egli, CP and Blum, JW 1998. Clinical, haematological, metabolic and endocrine traits during the first three months of life of suckling simmentaler calves held in a cow-calf operation. Zentralblatt für Veterinärmedizin. Reihe A 45, 99118.CrossRefGoogle Scholar
Farm Animal Welfare Committee (FAWC) 2015. Opinion on the welfare implications of nutritional management strategies for artificially-reared calves from birth to weaning. FAWC, University of Bristol, Bristol, UK.Google Scholar
Frieten, D, Gerbert, C, Koch, C, Dusel, G, Eder, K, Kanitz, E, Weitzel, JM and Hammon, HM 2017. Ad libitum milk replacer feeding, but not butyrate supplementation, affects growth performance as well as metabolic and endocrine traits in Holstein calves. Journal of Dairy Science 100, 66486661.CrossRefGoogle ScholarPubMed
Frieten, D, Gerbert, C, Koch, C, Dusel, G, Eder, K, Hoeflich, A, Mielenz, B and Hammon, HM 2018. Influence of ad libitum milk replacer feeding and butyrate supplementation on the systemic and hepatic insulin-like growth factor I and its binding proteins in Holstein calves. Journal of Dairy Science 101, 16611672.CrossRefGoogle ScholarPubMed
Geiger, AJ, Parsons, CLM, James, RE and Akers, RM 2016. Growth, intake, and health of Holstein heifer calves fed an enhanced preweaning diet with or without postweaning exogenous estrogen. Journal of Dairy Science 99, 39954004.CrossRefGoogle ScholarPubMed
Gelsinger, SL, Heinrichs, AJ and Jones, CM 2016. A meta-analysis of the effects of preweaned calf nutrition and growth on first-lactation performance. Journal of Dairy Science 99, 62066214.CrossRefGoogle ScholarPubMed
Georgiev, IP, Georgieva, TM, Pfaffl, M, Hammon, HM and Blum, JW 2003. Insulin-like growth factor and insulin receptors in intestinal mucosa of neonatal calves. Journal of Endocrinology 176, 121132.CrossRefGoogle ScholarPubMed
Gerbert, C, Frieten, D, Koch, C, Dusel, G, Eder, K, Stefaniak, T, Bajzert, J, Jawor, P, Tuchscherer, A and Hammon, HM 2018. Effects of ad libitum milk replacer feeding and butyrate supplementation on behavior, immune status, and health of Holstein calves in the postnatal period. Journal of Dairy Science 101, 73487360.CrossRefGoogle ScholarPubMed
Godden, S 2008. Colostrum management for dairy calves. Veterinary Clinics of North America: Food Animal Practice 24, 1939.Google ScholarPubMed
Gross, JJ and Bruckmaier, RM 2019. Invited review: metabolic challenges and adaptation during different functional stages of the mammary gland in dairy cows: perspectives for sustainable milk production. Journal of Dairy Science 102, 28282843.CrossRefGoogle ScholarPubMed
Gruse, J, Görs, S, Tuchscherer, A, Otten, W, Weitzel, JM, Metges, CC, Wolffram, S and Hammon, HM 2015. The effects of oral quercetin supplementation on splanchnic glucose metabolism in 1-week-old calves depend on diet after birth. Journal of Nutrition 145, 24862495.CrossRefGoogle ScholarPubMed
Guilloteau, P, Zabielski, R and Blum, JW 2009a. Gastrointestinal tract and digestion in the young ruminant: ontogenesis, adaptations, consequences and manipulations. Journal of Physiology and Pharmacology 60 (suppl. 3), 3746.Google Scholar
Guilloteau, P, Zabielski, R, Hammon, HM and Metges, CC 2009b. Adverse effects of nutritional programming during prenatal and early postnatal life, some aspects of regulation and potential prevention and treatments. Journal of Physiology and Pharmacology 60 (suppl. 3), 1735.Google ScholarPubMed
Hadorn, U, Hammon, H, Bruckmaier, RM and Blum, JW 1997. Delaying colostrum intake by one day has important effects on metabolic traits and on gastrointestinal and metabolic hormones in neonatal calves. Journal of Nutrition 127, 20112023.CrossRefGoogle ScholarPubMed
Haisan, J, Oba, M, Ambrose, DJ and Steele, MA 2018. Short communication: the effects of offering a high or low plane of milk preweaning on insulin-like growth factor and insulin-like growth factor binding proteins in dairy heifer calves. Journal of Dairy Science 101, 1144111446.CrossRefGoogle ScholarPubMed
Hammon, HM, Frieten, D, Gerbert, C, Koch, C, Dusel, G, Weikard, R and Kühn, C 2018. Different milk diets have substantial effects on the jejunal mucosal immune system of pre-weaning calves, as demonstrated by whole transcriptome sequencing. Scientific Reports 8, 1693.CrossRefGoogle ScholarPubMed
Hammon, HM, Schiessler, G, Nussbaum, A and Blum, JW 2002. Feed intake patterns, growth performance, and metabolic and endocrine traits in calves fed unlimited amounts of colostrum and milk by automate, starting in the neonatal period. Journal of Dairy Science 85, 33523362.CrossRefGoogle Scholar
Hammon, HM, Steinhoff-Wagner, J, Flor, J, Schönhusen, U and Metges, CC 2013. Lactation biology symposium: role of colostrum and colostrum components on glucose metabolism in neonatal calves. Journal of Animal Science 91, 685695.CrossRefGoogle ScholarPubMed
Hammon, HM, Steinhoff-Wagner, J, Schönhusen, U, Metges, CC and Blum, JW 2012. Energy metabolism in the newborn farm animal with emphasis on the calf: endocrine changes and responses to milk-born and systemic hormones. Domestic Animal Endocrinology 43, 171185.CrossRefGoogle ScholarPubMed
Hare, E, Norman, HD and Wright, JR 2006. Survival rates and productive herd life of dairy cattle in the United States. Journal of Dairy Science 89, 37133720.CrossRefGoogle ScholarPubMed
Huber, JT 1969. Development of the digestive and metabolic apparatus of the calf. Journal of Dairy Science 52, 13031315.CrossRefGoogle ScholarPubMed
Huber, K 2018. Invited review: resource allocation mismatch as pathway to disproportionate growth in farm animals–prerequisite for a disturbed health. Animal 12, 528536.CrossRefGoogle ScholarPubMed
Inabu, Y, Pyo, J, Pletts, S, Guan, LL, Steele, MA and Sugino, T 2019. Effect of extended colostrum feeding on plasma glucagon-like peptide-1 concentration in newborn calves. Journal of Dairy Science 102, 46194627.CrossRefGoogle ScholarPubMed
Ingvartsen, KL and Moyes, KM 2015. Factors contributing to immunosuppression in the dairy cow during the periparturient period. Japanese Journal of Veterinary Research 63 (suppl. 1), S1524.Google ScholarPubMed
Jacobi, SK and Odle, J 2012. Nutritional factors influencing intestinal health of the neonate. Advances in Nutrition 3, 687696.CrossRefGoogle ScholarPubMed
Jasper, J and Weary, DM 2002. Effects of ad libitum milk intake on dairy calves. Journal of Dairy Science 85, 30543058.CrossRefGoogle ScholarPubMed
Jensen, MB and Holm, L 2003. The effect of milk flow rate and milk allowance on feeding related behaviour in dairy calves fed by computer controlled milk feeders. Applied Animal Behaviour Science 82, 87100.CrossRefGoogle Scholar
Johnson, LR 1988. Regulation of gastrointestinal mucosal growth. Physiological Reviews 68, 456502.CrossRefGoogle ScholarPubMed
Kadowaki, T, Yamauchi, T, Kubota, N, Hara, K, Ueki, K and Tobe, K 2006. Adiponectin and adiponectin receptors in insulin resistance, diabetes, and the metabolic syndrome. Journal of Clinical Investigation 116, 17841792.CrossRefGoogle ScholarPubMed
Kenéz, Á, Koch, C, Korst, M, Kesser, J, Eder, K, Sauerwein, H and Huber, K 2018. Different milk feeding intensities during the first 4 weeks of rearing dairy calves: part 3: plasma metabolomics analysis reveals long-term metabolic imprinting in Holstein heifers. Journal of Dairy Science 101, 84468460.CrossRefGoogle ScholarPubMed
Kertz, AF, Hill, TM, Quigley, JD 3rd, Heinrichs, AJ, Linn, JG and Drackley, JK 2017. A 100-year review: calf nutrition and management. Journal of Dairy Science 100, 1015110172.CrossRefGoogle ScholarPubMed
Kesser, J, Hill, M, Heinz, JF, Koch, C, Rehage, J, Steinhoff-Wagner, J, Hammon, HM, Mielenz, B, Sauerwein, H and Sadri, H 2015. The rapid increase of circulating adiponectin in neonatal calves depends on colostrum intake. Journal of Dairy Science 98, 70447051.CrossRefGoogle ScholarPubMed
Khan, MA, Bach, A, Weary, DM and von Keyserlingk, MAG 2016. Invited review: transitioning from milk to solid feed in dairy heifers. Journal of Dairy Science 99, 885902.CrossRefGoogle ScholarPubMed
Khan, MA, Weary, DM and von Keyserlingk, MA 2011. Invited review: effects of milk ration on solid feed intake, weaning, and performance in dairy heifers. Journal of Dairy Science 94, 10711081.CrossRefGoogle ScholarPubMed
Khan, MA, Lee, HJ, Lee, WS, Kim, HS, Ki, KS, Hur, TY, Suh, GH, Kang, SJ and Choi, YJ 2007a. Structural growth, rumen development, and metabolic and immune responses of Holstein male calves fed milk through step-down and conventional methods. Journal of Dairy Science 90, 33763387.CrossRefGoogle ScholarPubMed
Khan, MA, Lee, HJ, Lee, WS, Kim, HS, Kim, SB, Ki, KS, Ha, JK, Lee, HG and Choi, YJ 2007b. Pre- and postweaning performance of Holstein female calves fed milk through step-down and conventional methods. Journal of Dairy Science 90, 876885.CrossRefGoogle ScholarPubMed
Koch, C, Gerbert, C, Frieten, D, Dusel, G, Eder, K, Zitnan, R and Hammon, HM 2019. Effects of ad libitum milk replacer feeding and butyrate supplementation on the epithelial growth and development of the gastrointestinal tract in Holstein calves. Journal of Dairy Science 102, 85138526.CrossRefGoogle ScholarPubMed
Korst, M, Koch, C, Kesser, J, Müller, U, Romberg, FJ, Rehage, J, Eder, K and Sauerwein, H 2017. Different milk feeding intensities during the first 4 weeks of rearing in dairy calves: part 1: effects on performance and production from birth over the first lactation. Journal of Dairy Science 100, 30963108.CrossRefGoogle ScholarPubMed
Langel, SN, Wark, WA, Garst, SN, James, RE, McGilliard, ML, Petersson-Wolfe, CS and Kanevsky-Mullarky, I 2015. Effect of feeding whole compared with cell-free colostrum on calf immune status: the neonatal period. Journal of Dairy Science 98, 37293740.CrossRefGoogle ScholarPubMed
Leal, LN, Romao, JM, Hooiveld, GJ, Soberon, F, Berends, H, Boekshoten, MV, Van Amburgh, ME, Martin-Tereso, J and Steele, MA 2018. Nutrient supply alters transcriptome regulation in adipose tissue of pre-weaning Holstein calves. PLoS ONE 13, e0201929.CrossRefGoogle ScholarPubMed
Lepine, AJ, Boyd, RD and Whitehead, DM 1991. Effect of colostrum intake on hepatic gluconeogenesis and fatty acid oxidation in the neonatal pig. Journal of Animal Science 69, 19661974.CrossRefGoogle ScholarPubMed
Liebler-Tenorio, EM, Riedel-Caspari, G and Pohlenz, JF 2002. Uptake of colostral leukocytes in the intestinal tract of newborn calves. Veterinary Immunology and Immunopathology 85, 3340.CrossRefGoogle ScholarPubMed
Maccari, P, Wiedemann, S, Kunz, HJ, Piechotta, M, Sanftleben, P and Kaske, M 2015. Effects of two different rearing protocols for Holstein bull calves in the first 3 weeks of life on health status, metabolism and subsequent performance. Journal of Animal Physiology and Animal Nutrition 99, 737746.CrossRefGoogle ScholarPubMed
MacPherson, J, Meale, SJ, Macmillan, K, Haisan, J, Bench, CJ, Oba, M and Steele, MA 2019. Effects of feeding frequency of an elevated plane of milk replacer and calf age on behavior, and glucose and insulin kinetics in male Holstein calves. Animal 13, 13851393.CrossRefGoogle ScholarPubMed
Malmuthuge, N and Guan, LL 2017. Understanding the gut microbiome of dairy calves: opportunities to improve early-life gut health. Journal of Dairy Science 100, 59966005.CrossRefGoogle ScholarPubMed
Mathers, JC 1998. Nutrient regulation of intestinal proliferation and apoptosis. Proceedings of the Nutrition Society 57, 219223.CrossRefGoogle ScholarPubMed
Mee, JF 2008. Newborn dairy calf management. Veterinary Clinics of North America: Food Animal Practice 24, 117.Google ScholarPubMed
Miller-Cushon, EK, Bergeron, R, Leslie, KE and DeVries, TJ 2013. Effect of milk feeding level on development of feeding behavior in dairy calves. Journal of Dairy Science 96, 551564.CrossRefGoogle ScholarPubMed
Miller-Cushon, EK and DeVries, TJ 2015. Invited review: development and expression of dairy calf feeding behaviour. Canadian Journal of Animal Science 95, 341350.CrossRefGoogle Scholar
Molès, JP, Tuaillon, E, Kankasa, C, Bedin, AS, Nagot, N, Marchant, A, McDermid, JM and Van de Perre, P 2018. Breastmilk cell trafficking induces microchimerism-mediated immune system maturation in the infant. Pediatric Allergy and Immunology 29, 133143.CrossRefGoogle ScholarPubMed
Mylonas, PG, Matsouka, PT, Papandoniou, EV, Vagianos, C, Kalfarentzos, F and Alexandrides, TK 2000. Growth hormone and insulin-like growth factor I protect intestinal cells from radiation induced apoptosis. Molecular and Cellular Endocrinology 160, 115122.CrossRefGoogle ScholarPubMed
Naeem, A, Drackley, JK, Stamey, J and Loor, JJ 2012. Role of metabolic and cellular proliferation genes in ruminal development in response to enhanced plane of nutrition in neonatal Holstein calves. Journal of Dairy Science 95, 18071820.CrossRefGoogle ScholarPubMed
Naeem, A, Drackley, JK, Lanier, JS, Everts, RE, Rodriguez-Zas, SL and Loor, JJ 2014. Ruminal epithelium transcriptome dynamics in response to plane of nutrition and age in young Holstein calves. Functional & Integrative Genomics 14, 261273.CrossRefGoogle ScholarPubMed
Nissen, A, Andersen, PH, Bendixen, E, Ingvartsen, KL and Rontved, CM 2017. Colostrum and milk protein rankings and ratios of importance to neonatal calf health using a proteomics approach. Journal of Dairy Science 100, 27112728.CrossRefGoogle ScholarPubMed
Nonnecke, BJ, Foote, MR, Smith, JM, Pesch, BA and Van Amburgh, ME 2003. Composition and functional capacity of blood mononuclear leukocyte populations from neonatal calves on standard and intensified milk replacer diets. Journal of Dairy Science 86, 35923604.CrossRefGoogle ScholarPubMed
OIE 2017. Terrestrial Animal Health Code. Section 7. Animal Welfare. Retrieved on 3 July 2019 from http://oie.int/international-standard-setting/terrestrial-codeGoogle Scholar
Ollivett, TL, Nydam, DV, Linden, TC, Bowman, DD and Van Amburgh, ME 2012. Effect of nutritional plane on health and performance in dairy calves after experimental infection with Cryptosporidium parvum. Journal of the American Veterinary Medical Association 241, 15141520.CrossRefGoogle ScholarPubMed
Ontsouka, EC, Albrecht, C and Bruckmaier, RM 2016. Invited review: growth-promoting effects of colostrum in calves based on interaction with intestinal cell surface receptors and receptor-like transporters. Journal of Dairy Science 99, 41114123.CrossRefGoogle ScholarPubMed
Probo, M, Pascottini, OB, LeBlanc, S, Opsomer, G and Hostens, M 2018. Association between metabolic diseases and the culling risk of high-yielding dairy cows in a transition management facility using survival and decision tree analysis. Journal of Dairy Science 101, 94199429.CrossRefGoogle Scholar
Prokop, L, Kaske, M, Maccari, P, Lucius, R, Kunz, HJ and Wiedemann, S 2015. Intensive rearing of male calves during the first three weeks of life has long-term effects on number of islets of Langerhans and insulin stained area in the pancreas. Journal of Animal Science 93, 988998.CrossRefGoogle ScholarPubMed
Qi, Y, Zhao, X, Huang, D, Pan, X, Yang, Y, Zhao, H, Hu, H and Cheng, G 2018. Exploration of the relationship between intestinal colostrum or milk, and serum metabolites in neonatal calves by metabolomics analysis. Journal of Agricultural and Food Chemistry 66, 72007208.CrossRefGoogle ScholarPubMed
Ramachandran, A, Madesh, M and Balasubramanian, KA 2000. Apoptosis in the intestinal epithelium: its relevance in normal and pathophysiological conditions. Journal of Gastroenterology and Hepatology 15, 109120.CrossRefGoogle ScholarPubMed
Renaville, R, Van Eenaeme, C, Breier, BH, Vleurick, L, Bertozzi, C, Gengler, N, Hornick, JL, Parmentier, I, Istasse, L, Haezebroeck, V, Massart, S and Portetelle, D 2000. Feed restriction in young bulls alters the onset of puberty in relationship with plasma insulin-like growth factor-I (IGF-I) and IGF-binding proteins. Domestic Animal Endocrinology 18, 165176.CrossRefGoogle ScholarPubMed
Roffler, B, Fäh, A, Sauter, SN, Hammon, HM, Gallmann, P, Brem, G and Blum, JW 2003. Intestinal morphology, epithelial cell proliferation, and absorptive capacity in neonatal calves fed milk-born insulin-like growth factor-I or a colostrum extract. Journal of Dairy Science 86, 17971806.CrossRefGoogle ScholarPubMed
Rosenberger, K, Costa, JHC, Neave, HW, von Keyserlingk, MAG and Weary, DM 2017. The effect of milk allowance on behavior and weight gains in dairy calves. Journal of Dairy Science 100, 504512.CrossRefGoogle ScholarPubMed
Sadri, H, Steinhoff-Wagner, J, Hammon, HM, Bruckmaier, RM, Gors, S and Sauerwein, H 2017. Mammalian target of rapamycin signaling and ubiquitin proteasome-related gene expression in 3 different skeletal muscles of colostrum- versus formula-fed calves. Journal of Dairy Science 100, 94289441.CrossRefGoogle ScholarPubMed
Santos, A, Giraldez, FJ, Frutos, J and Andres, S 2019. Liver transcriptomic and proteomic profiles of preweaning lambs are modified by milk replacer restriction. Journal of Dairy Science 102, 11941204.CrossRefGoogle ScholarPubMed
Sauter, SN, Roffler, B, Philipona, C, Morel, C, Rome, V, Guilloteau, P, Blum, JW and Hammon, HM 2004. Intestinal development in neonatal calves: effects of glucocorticoids and dependence of colostrum feeding. Biology of the Neonate 85, 94104.CrossRefGoogle ScholarPubMed
Schäff, CT, Gruse, J, Maciej, J, Mielenz, M, Wirthgen, E, Hoeflich, A, Schmicke, M, Pfuhl, R, Jawor, P, Stefaniak, T and Hammon, HM 2016. Effects of feeding milk replacer ad libitum or in restricted amounts for the first five weeks of life on the growth, metabolic adaptation, and immune status of newborn calves. PLoS ONE 11, e0168974.CrossRefGoogle ScholarPubMed
Schäff, CT, Gruse, J, Maciej, J, Pfuhl, R, Zitnan, R, Rajsky, M and Hammon, HM 2018. Effects of feeding unlimited amounts of milk replacer for the first 5 weeks of age on rumen and small intestinal growth and development in dairy calves. Journal of Dairy Science 101, 783793.CrossRefGoogle ScholarPubMed
Schiessler, G, Nussbaum, A, Hammon, HM and Blum, JW 2002. Calves sucking, colostrum and milk from their dams or from an automatic feeding station starting in the neonatal period: metabolic and endocrine traits and growth performance. Animal Science 74, 431444.CrossRefGoogle Scholar
Smith, JM, Van Amburgh, ME, Diaz, MC, Lucy, MC and Bauman, DE 2002. Effect of nutrient intake on the development of the somatotropic axis and its responsiveness to GH in Holstein bull calves. Journal of Animal Science 80, 15281537.CrossRefGoogle ScholarPubMed
Soberon, F and Van Amburgh, ME 2017. Effects of preweaning nutrient intake in the developing mammary parenchymal tissue. Journal of Dairy Science 100, 49965004.CrossRefGoogle ScholarPubMed
Steele, MA, Penner, GB, Chaucheyras-Durand, F and Guan, LL 2016. Development and physiology of the rumen and the lower gut: targets for improving gut health. Journal of Dairy Science 99, 49554966.CrossRefGoogle ScholarPubMed
Steinhoff-Wagner, J, Gors, S, Junghans, P, Bruckmaier, RM, Kanitz, E, Metges, CC and Hammon, HM 2011. Intestinal glucose absorption but not endogenous glucose production differs between colostrum- and formula-fed neonatal calves. Journal of Nutrition 141, 4855.CrossRefGoogle Scholar
Steinhoff-Wagner, J, Zitnan, R, Schönhusen, U, Pfannkuche, H, Hudakova, M, Metges, CC and Hammon, HM 2014. Diet effects on glucose absorption in the small intestine of neonatal calves: importance of intestinal mucosal growth, lactase activity, and glucose transporters. Journal of Dairy Science 97, 63586369.CrossRefGoogle ScholarPubMed
Stelwagen, K, Carpenter, E, Haigh, B, Hodgkinson, A and Wheeler, TT 2009. Immune components of bovine colostrum and milk. Journal of Animal Science 87, 39.CrossRefGoogle ScholarPubMed
Tautenhahn, A. 2017. Risikofaktoren für eine erhöhte Kälbersterblichkeit und geringe Tageszunahmen von Aufzuchtkälbern in norddeutschen Milchkuhhaltungen. Dissertation for Doctor of Veterinary Medicine. Free University of Berlin, Berlin, Germany.Google Scholar
Urie, NJ, Lombard, JE, Shivley, CB, Kopral, CA, Adams, AE, Earleywine, TJ, Olson, JD and Garry, FB 2018. Preweaned heifer management on US dairy operations: part V. Factors associated with morbidity and mortality in preweaned dairy heifer calves. Journal of Dairy Science 101, 92299244.CrossRefGoogle ScholarPubMed
Van Amburgh, ME and Soberon, F 2013. The role of calf nutrition and management on lifetime productivity of dairy cattle. In Proceedings of the Cow Longevity Conference, 28–29 August 2013, Hamra Farm, Tumba, Sweden, pp. 178197.Google Scholar
Von Keyserlingk, MAG, Rushen, J, de Passille, AM and Weary, DM 2009. The welfare of dairy cattle–key concepts and the role of science. Journal of Dairy Science 92, 41014111.CrossRefGoogle Scholar
Wang, P, Drackley, JK, Stamey-Lanier, JA, Keisler, D and Loor, JJ 2014. Effects of level of nutrient intake and age on mammalian target of rapamycin, insulin, and insulin-like growth factor-1 gene network expression in skeletal muscle of young Holstein calves. Journal of Dairy Science 97, 383391.CrossRefGoogle ScholarPubMed
Weikard, R, Hadlich, F, Hammon, HM, Frieten, D, Gerbert, C, Koch, C, Dusel, G and Kuehn, C 2018. Long noncoding RNAs are associated with metabolic and cellular processes in the jejunum mucosa of pre-weaning calves in response to different diets. Oncotarget 9, 2105221069.CrossRefGoogle ScholarPubMed
Welboren, AC, Leal, LN, Steele, MA, Khan, MA and Martin-Tereso, J 2019. Performance of ad libitum fed dairy calves weaned using fixed and individual methods. Animal 13, 18911898.CrossRefGoogle ScholarPubMed
Xu, RJ 1996. Development of the newborn GI tract and its relation to colostrum milk intake: a review. Reproduction Fertility and Development 8, 3548.CrossRefGoogle ScholarPubMed
Zhao, XW, Qi, YX, Huang, DW, Pan, XC, Cheng, GL, Zhao, HL and Yang, YX 2018. Changes in serum metabolites in response to ingested colostrum and milk in neonatal calves, measured by nuclear magnetic resonance-based metabolomics analysis. Journal of Dairy Science 101, 71687181.CrossRefGoogle ScholarPubMed
Figure 0

Table 1 Effects of intensive milk feeding immediately after birth on growth, development, behaviour and immune response in preweaning calves1