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To optimize first lactation and lifetime milk yield, growth benchmarks were established to help meet the appropriate growth objectives of breeding weight and age at an economically viable time and to achieve the optimum body size and composition at first calving. These guidelines provide a framework that helps to minimize overfeeding and, thus, potential overconditioning of heifers, which can lead to postpartum metabolic issues and reduced milk yield. Concerns still exist that mammary development is impaired when body weight gain exceeds a certain threshold, which would negatively affects milk yield. The objective of this review was to integrate concepts of nutrient requirements, body growth and composition, mammary development, and milk yield to provide a systems-based perspective on first-lactation milk differences that have been associated with mammary development. Work in the early 1980s described the effect of high energy intake on mammary development and the relationship with circulating growth hormone linked the relationship between prepubertal growth, mammary development, and future milk yield. The primary outcome of that research was to provide an intuitive mechanism to explain why rapid growth during the prepubertal phase resulted in reduced milk yield. The observation of reduced mammary development could be repeated in almost every experiment, leading to the conclusion that high energy intake and increased average daily gain reduced mammary development through altered hormone status or some signaling processes. However, further work that looked at mammary development over the entire prepubertal growth phase recognized that mammary development was not reduced by high energy intake, and instead accumulated at a constant rate; thus, overall mammary parenchymal growth was a function of the time to reach puberty and the associated signals to change from allometric mammary growth. The mammary gland, similar to most reproductive organs, grows in proportion to the size of the body and not in proportion to nutrient intake during the postweaning, prepubertal phase. First-lactation milk yield, mammary development, and body composition will be further discussed in the context of mechanisms and opportunities., (Copyright © 2019 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

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., (The Authors. Published by FASS Inc. and Elsevier Inc. on behalf of the American Dairy Science Association®. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).)

Data from calf studies conducted over the past 20 yr has shown that preweaning nutrient intake, from milk or milk replacer, can have profound effects on development of the calf that enhance first lactation and lifetime productivity. Many of the studies show positive but not significant effects of preweaning nutrition on long-term productivity, primarily due to a lack of power, usually due to inadequate animal numbers per treatment. Meta-analyses were conducted using Comprehensive Meta-Analysis (CMA) software (version 2.2.064; Biostat, Englewood, NJ; Borenstein et al., 2005) to evaluate the effects of preweaning nutrient intake and preweaning ADG on first lactation milk production from studies where milk yield and preweaning treatment data were adequately described. Currently, 12 studies have been reported, describing milk yield of calves that had been fed various levels of preweaning nutrients, from both milk and milk replacer, and 11 evaluated the effect of preweaning ADG on long-term productivity. The estimated effect size for treatment (level of milk or milk replacer intake) and ADG were calculated. Using a random effects model, the overall milk yield response based on treatment was 435 ± 117 kg/lactation (P < 0.001), demonstrating that among the data sets milk yield in the first lactation was increased by increasing nutrient intake from milk or milk replacer in preweaned calves. Meta-regression of the effect of ADG resulted in the following equation: milk yield = -106 kg + 1,551.4 kg × ADG (kg/d; P = 0.01) indicating that for every kilogram of preweaning ADG, first lactation milk yield increased by 1,550 kg. Furthermore, the meta-analysis yielded an odds ratio of 2.09 (P = 0.001) indicating that calves fed for greater preweaning ADG were 2 times more likely to have greater milk yield in the first lactation. Finally, the 2001 Dairy NRC calf model was used to estimate the difference, within study, of intake over maintenance from milk replacer, using either the actual data or the published ADG; the difference in intake over maintenance was used as a predictor in a meta-regression and resulted in the following equation: milk yield = -60 kg + 1,100 kg × estimated difference in intake over maintenance (P = 0.02). The data demonstrate that nutrient intake from milk or milk replacer during the preweaning period positively impacted long-term productivity of dairy calves and provides new management opportunities to improve milk yield of dairy cattle. These data appear to indicate that there are significant developmental functions being programmed in the neonatal calf that require further investigation.

The preweaning management of dairy calves over the last 30 yr has focused on mortality, early weaning, and rumen development. Recent studies suggest that nutrient intake from milk or milk replacer during the preweaning period alters the phenotypic expression for milk yield. The objective of this study was to investigate the relationship between nutrient intake from milk replacer and pre- and postweaning growth rate with lactation performance in the Cornell dairy herd and a commercial dairy farm. The analysis was conducted using traditional 305-d first-lactation milk yield and residual lactation yield estimates from a test-day model (TDM) to analyze the lactation records over multiple lactations. The overall objective of the calf nutrition program in both herds was to double the birth weight of calves by weaning through increased milk replacer and starter intake. First-lactation 305-d milk yield and residuals from the TDM were generated from 1,244 and 624 heifers from the Cornell herd and from the commercial farm, respectively. The TDM was used to generate lactation residuals after accounting for the effects of test day, calving season, days in milk, days pregnant, lactation number, and year. In addition, lactation residuals were generated for cattle with multiple lactations to determine if the effect of preweaning nutrition could be associated with lifetime milk yield. Factors such as preweaning average daily gain (ADG), energy intake from milk replacer as a multiple of maintenance, and other growth outcomes and management variables were regressed on TDM milk yield data. In the Cornell herd, preweaning ADG, ranged from 0.10 to 1.58 kg, and was significantly correlated with first-lactation yield; for every 1 kg of preweaning ADG, heifers, on average, produced 850 kg more milk during their first lactation and 235 kg more milk for every Mcal of metabolizable energy intake above maintenance. In the commercial herd, for every 1 kg of preweaning ADG, milk yield increased by 1,113 kg in the first lactation and further, every 1 kg of prepubertal ADG was associated with a 3,281 kg increase in first-lactation milk yield. Among the 2 herds, preweaning ADG accounted for 22% of the variation in first-lactation milk yield as analyzed with the TDM. These results indicate that increased growth rate before weaning results in some form of epigenetic programming that is yet to be understood, but has positive effects on lactation milk yield. This analysis identifies nutrition and management of the preweaned calf as major environmental factors influencing the expression of the genetic capacity of the animal for milk yield., (Copyright © 2012 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)

Results of previous studies have shown that increased milking frequency (IMF) during early lactation results in increased milk yield not only during the period of IMF but also after cows have returned to a decreased milking frequency. The cellular mechanisms underpinning this increased milk yield and the overall effects of IMF on metabolism have not been well characterized. The objective of this study was to determine the effects of IMF on metabolism and mammary epithelial cell proliferation in dairy cows. Thirty primiparous and 30 multiparous Holstein cows were assigned randomly at calving to 1 of 2 treatments. The control group was milked twice daily (2x) for 119 d, whereas the IMF group was milked 4 times daily (4x) from d 2 postcalving until d 21 and then 2x from d 22 until d 119. Overall milk yield did not differ between treatments throughout the 119 d monitored; however, the interaction of treatment by week was significant in that IMF cows yielded 4.8kg/d more milk than control cows during wk 2 and 3 and had similar levels of milk yield during the remainder of the study period. Reanalysis of data excluding data from cows subjected to mammary biopsy suggested that the mammary biopsy procedure contributed to the lack of overall responses of milk yield, but that responses overall to IMF were greater in primiparous cows compared with multiparous cows. Plasma nonesterified fatty acid concentrations were elevated in multiparous cows subjected to IMF during the period of IMF, but were not influenced by treatment in primiparous cows. Plasma beta-hydroxybutyrate concentrations were not affected by treatment. Mammary tissue was collected by biopsy in a subset of cows (n=8 cows per parity and treatment) at calving and at d 21 and 75 postpartum and used for immunohistochemical localization of the cell proliferation antigen, Ki67. Effects of treatment on mammary epithelial cell proliferation were not significant, suggesting that other mechanisms must be responsible for carryover effects of IMF on lactational performance., (Copyright 2010 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.)