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Review Article|Articles in Press

INVITED REVIEW: Nutritional and management factors that influence colostrum production and composition in dairy cows

Open AccessPublished:January 19, 2024DOI:https://doi.org/10.3168/jds.2023-24349

      ABSTRACT

      Colostrum is a rich source of nutritional and non-nutritional components and is recognized as essential to transfer passive immunity to newborn calves. Because of the individual and seasonal variability in colostrum yield and composition, maintaining an adequate supply of high-quality colostrum year-round remains a challenge for commercial dairy producers. In this narrative review, we described the individual, seasonal, and herd-level variability of colostrum production and summarized the association between individual animal factors such as parity, sex of the calf, calf birth weight, as well as indicators of the cow's metabolic status and the yield and composition of colostrum. Further, we reviewed the current knowledge on the influence of prepartum nutrition and management strategies on colostrum production. Research on the metabolizable energy and protein supplied in the prepartum diet as well as into the inclusion and source of vitamins, minerals, and feed additives suggests prepartum nutrition influences the yield, quality, and composition of colostrum. Furthermore, the prepartum environment and dry period length remain influential factors in the production of colostrum. However, additional research is needed to understand the mechanisms by which prepartum nutrition and management affects colostrum production. Lastly, time to colostrum harvest and oxytocin administration as well as the current knowledge on the effect of heat-treatment and colostrum storage strategies on colostral components were discussed. To conclude, we identify critical gaps in knowledge for future focus of investigation in colostrum research.

      Key Words

      INTERPRETIVE SUMMARY Maintaining an adequate supply of high-quality colostrum is essential for rearing healthy calves but remains a challenge because of the individual and seasonal variability in colostrum production. This narrative review provides an overview of the variability in colostrum production and discusses the current knowledge on the effect of prepartum nutritional and managerial factors on colostrum production. Moreover, we discuss the effect of post-harvest practices on colostral components and identify critical gaps in knowledge for future focus of investigation.

      INTRODUCTION

      Mounting evidence supports the importance of colostrum for raising healthy calves (
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      Impact of 2 versus 1 colostrum meals on failure of transfer of passive immunity, pre-weaning morbidity and mortality, and performance of dairy calves in a large dairy herd.
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      Comparison of calf morbidity, mortality, and future performance across categories of passive immunity: A retrospective cohort study in a dairy herd.
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      Association between transfer of passive immunity, health, and performance of female dairy calves from birth to weaning.
      ). Because the cotyledonary synepitheliochorial bovine placenta inhibits transfer of maternal antibodies into fetal circulation, newborn dairy calves rely on timely ingestion of high-quality colostrum with low bacterial contamination to transfer passive immunity (TPI) as well as for nutrients and other bioactive components (
      • Fischer-Tlustos A.J.
      • Cabus A.J.L.
      • Hare K.S.
      • Wood K.
      • Steele M.
      Invited Review: Effects of colostrum management on transfer of passive immunity and the potential role of colostral bioactive components on neonatal calf development and metabolism.
      ,
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      • Heinrichs A.J.
      Invited review: The importance of colostrum in the newborn dairy calf.
      ). Poor TPI in calves has been associated with lower average daily gain, a greater risk for preweaning morbidity and mortality, and a lower likelihood to reach first insemination and calving (
      • Crannell P.
      • Abuelo A.
      Comparison of calf morbidity, mortality, and future performance across categories of passive immunity: A retrospective cohort study in a dairy herd.
      ,
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      • Heuwieser W.
      • Borchardt S.
      Association between transfer of passive immunity, health, and performance of female dairy calves from birth to weaning.
      ). Further, colostrum intake aids in the development of the gastrointestinal tract (
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      Lactation Biology Symposium: role of colostrum and colostrum components on glucose metabolism in neonatal calves.
      ,
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      Heat-treated colostrum feeding promotes beneficial bacteria colonization in the small intestine of neonatal calves.
      ,
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      Importance of colostrum supply and milk feeding intensity on gastrointestinal and systemic development in calves.
      ,
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      • Hare K.
      • Pletts S.
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      • Haines D.
      • Sugino T.
      • Guan L.
      • Steele M.
      Feeding colostrum or a 1: 1 colostrum: milk mixture for 3 days postnatal increases small intestinal development and minimally influences plasma glucagon-like peptide-2 and serum insulin-like growth factor-1 concentrations in Holstein bull calves.
      ) and has been shown to positively affect first lactation milk production (
      • Faber S.
      • Faber N.
      • McCauley T.
      • Ax R.
      Case study: effects of colostrum ingestion on lactational performance.
      ,
      • Abuelo A.
      • Cullens F.
      • Hanes A.
      • Brester J.L.
      Impact of 2 versus 1 colostrum meals on failure of transfer of passive immunity, pre-weaning morbidity and mortality, and performance of dairy calves in a large dairy herd.
      ). As such, it is recommended to feed calves 8.5 to 10% of BW (
      • Conneely M.
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      • Doherty M.L.
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      Effect of feeding colostrum at different volumes and subsequent number of transition milk feeds on the serum immunoglobulin G concentration and health status of dairy calves.
      , Godden et al., 2019) of high-quality (≥50 g of IgG/L; ≥ 22.0% Brix) colostrum with low bacterial contamination (TPC < 100,000 cfu/mL; fecal coliforms < 10,000 cfu/mL;
      • McGuirk S.M.
      • Collins M.
      Managing the production, storage, and delivery of colostrum.
      ) at birth. A growing number of producers are also implementing an extended colostrum (
      • Kehoe S.I.
      • Jayarao B.M.
      • Heinrichs A.J.
      A survey of bovine colostrum composition and colostrum management practices on Pennsylvania dairy farms.
      ,
      • Robbers L.
      • Bijkerk H.J.C.
      • Koets A.P.
      • Benedictus L.
      • Nielen M.
      • Jorritsma R.
      Survey on Colostrum Management by Dairy Farmers in the Netherlands.
      ,
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      ) or transition milk feeding program to promote preweaning health, growth, and development (
      • Conneely M.
      • Berry D.P.
      • Murphy J.P.
      • Lorenz I.
      • Doherty M.L.
      • Kennedy E.
      Effect of feeding colostrum at different volumes and subsequent number of transition milk feeds on the serum immunoglobulin G concentration and health status of dairy calves.
      ,
      • Kargar S.
      • Roshan M.
      • Ghoreishi S.
      • Akhlaghi A.
      • Kanani M.
      • Shams-Abadi A.A.
      • Ghaffari M.
      Extended colostrum feeding for 2 weeks improves growth performance and reduces the susceptibility to diarrhea and pneumonia in neonatal Holstein dairy calves.
      ,
      • Pyo J.
      • Hare K.
      • Pletts S.
      • Inabu Y.
      • Haines D.
      • Sugino T.
      • Guan L.
      • Steele M.
      Feeding colostrum or a 1: 1 colostrum: milk mixture for 3 days postnatal increases small intestinal development and minimally influences plasma glucagon-like peptide-2 and serum insulin-like growth factor-1 concentrations in Holstein bull calves.
      ,
      • Van Soest B.
      • Cullens F.
      • VandeHaar M.
      • Nielsen M.W.
      Effects of transition milk and milk replacer supplemented with colostrum replacer on growth and health of dairy calves.
      ,
      • Abuelo A.
      • Cullens F.
      • Hanes A.
      • Brester J.L.
      Impact of 2 versus 1 colostrum meals on failure of transfer of passive immunity, pre-weaning morbidity and mortality, and performance of dairy calves in a large dairy herd.
      ) as well as selling colostrum for production of commercial products (
      • Costa A.
      • Sneddon N.
      • Goi A.
      • Visentin G.
      • Mammi L.
      • Savarino E.
      • Zingone F.
      • Formigoni A.
      • Penasa M.
      • De Marchi M.
      Invited review: Bovine colostrum, a promising ingredient for humans and animals—Properties, processing technologies, and uses.
      ,
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      ). Thus, an adequate supply of high-quality colostrum is a critical component for the viability of commercial dairy farms.
      Research on the epidemiology of colostrum production has exposed the individual and seasonal variability (
      • Gavin K.
      • Neibergs H.
      • Hoffman A.
      • Kiser J.N.
      • Cornmesser M.A.
      • Haredasht S.A.
      • Martinez-Lopez B.
      • Wenz J.R.
      • Moore D.A.
      Low colostrum yield in Jersey cattle and potential risk factors.
      ,
      • Borchardt S.
      • Sutter F.
      • Heuwieser W.
      • Venjakob P.
      Management-related factors in dry cows and their associations with colostrum quantity and quality on a large commercial dairy farm.
      ,
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      ) and prompted investigations to discover animal, environmental, nutritional, and managerial factors that influence the yield as well as IgG and component concentrations of colostrum. Further, dairy producers put forth equipment and labor resources required to harvest, store, reduce contamination, and feed high-quality colostrum. As such, this narrative review will describe the variability in colostrum production, discuss prepartum nutritional and management strategies as well as post-harvest practices associated with the production of colostrum and the preservation of colostral components, and identify knowledge gaps to direct future focus of investigation. The keywords “colostrum” and “cow” as well as other relevant keywords were used to identify literature that pertained to each variable discussed in this narrative review. Additional references were identified through citation mining. Unless otherwise stated, we refer to colostrum as the first milking only and to data from Holstein dairy cattle.

      VARIABILITY IN COLOSTRUM PRODUCTION

      Seasonality

      Of interest, colostrum yield (
      • Karl M.
      • Staufenbiel R.
      Factors influencing the amount of initial colostrum in Holstein-Friesian cows and their relationship to postpartum calcium concentration.
      ,
      • Gavin K.
      • Neibergs H.
      • Hoffman A.
      • Kiser J.N.
      • Cornmesser M.A.
      • Haredasht S.A.
      • Martinez-Lopez B.
      • Wenz J.R.
      • Moore D.A.
      Low colostrum yield in Jersey cattle and potential risk factors.
      ,
      • Borchardt S.
      • Sutter F.
      • Heuwieser W.
      • Venjakob P.
      Management-related factors in dry cows and their associations with colostrum quantity and quality on a large commercial dairy farm.
      ), Brix % or IgG concentration (
      • Zarei S.
      • Ghorbani G.R.
      • Khorvash M.
      • Martin O.B.
      • Mahdavi A.H.
      • Riasi A.
      The impact of season, parity, and volume of colostrum on Holstein dairy cows colostrum composition.
      ,
      • Borchardt S.
      • Sutter F.
      • Heuwieser W.
      • Venjakob P.
      Management-related factors in dry cows and their associations with colostrum quantity and quality on a large commercial dairy farm.
      ), and composition (
      • Zarei S.
      • Ghorbani G.R.
      • Khorvash M.
      • Martin O.B.
      • Mahdavi A.H.
      • Riasi A.
      The impact of season, parity, and volume of colostrum on Holstein dairy cows colostrum composition.
      ,
      • Soufleri A.
      • Banos G.
      • Panousis N.
      • Fletouris D.
      • Arsenos G.
      • Kougioumtzis A.
      • Valergakis G.E.
      Evaluation of Factors Affecting Colostrum Quality and Quantity in Holstein Dairy Cattle.
      ) exhibit seasonality. Colostrum yield was greatest in June (6.6 kg) and decreased to its lowest average yield of 1.3 kg in December in multiparous Jersey cows from a single dairy located in Texas, United States (
      • Gavin K.
      • Neibergs H.
      • Hoffman A.
      • Kiser J.N.
      • Cornmesser M.A.
      • Haredasht S.A.
      • Martinez-Lopez B.
      • Wenz J.R.
      • Moore D.A.
      Low colostrum yield in Jersey cattle and potential risk factors.
      ). Primiparous Jersey cows also experience seasonality in colostrum yield (June: 6.5 kg; December: 4.2 kg), albeit less than multiparous cows (
      • Gavin K.
      • Neibergs H.
      • Hoffman A.
      • Kiser J.N.
      • Cornmesser M.A.
      • Haredasht S.A.
      • Martinez-Lopez B.
      • Wenz J.R.
      • Moore D.A.
      Low colostrum yield in Jersey cattle and potential risk factors.
      ). In a study conducted in Germany, average colostrum yield from primiparous Holstein cows was greatest in April (4.1 ± 0.3 kg) and lowest during November (3.2 ± 0.3 kg) and average colostrum yield from multiparous cows peaked in May (5.5 ± 0.3 kg) and was lowest in October (3.8 ± 0.3 kg), respectively (
      • Borchardt S.
      • Sutter F.
      • Heuwieser W.
      • Venjakob P.
      Management-related factors in dry cows and their associations with colostrum quantity and quality on a large commercial dairy farm.
      ). Conversely, authors in the United States have associated calving during the summer (June–August) with reduced colostrum IgG concentration (
      • Morin D.E.
      • Constable P.D.
      • Maunsell F.P.
      • McCoy G.C.
      Factors associated with colostral specific gravity in dairy cows.
      ,
      • Rossi R.M.
      • Cullens F.M.
      • Bacigalupo P.
      • Sordillo L.M.
      • Abuelo A.
      Changes in biomarkers of metabolic stress during late gestation of dairy cows associated with colostrum volume and immunoglobulin content.
      ). Seasonality was also observed for other colostral components in samples collected in Northern Greece with greatest concentrations of fat in the spring (March–May), protein in the fall (September–November) and winter (December–February), and lactose in the fall, winter, and spring (
      • Soufleri A.
      • Banos G.
      • Panousis N.
      • Fletouris D.
      • Arsenos G.
      • Kougioumtzis A.
      • Valergakis G.E.
      Evaluation of Factors Affecting Colostrum Quality and Quantity in Holstein Dairy Cattle.
      ) and these associations might be related to changes in light as well as temperature and humidity exposure (discussed below). Although the inverse seasonal relationship between yield and IgG concentration has been hypothesized to be influenced by IgG dilution (
      • Pritchett L.C.
      • Gay C.C.
      • Besser T.E.
      • Hancock D.D.
      Management and production factors influencing immunoglobulin G1 concentration in colostrum from Holstein cows.
      ,
      • Guy M.A.
      • McFadden T.B.
      • Cockrell D.C.
      • Besser T.E.
      Regulation of colostrum formation in beef and dairy cows.
      ,
      • Kehoe S.
      • Heinrichs A.J.
      • Moody M.
      • Jones C.
      • Long M.
      Comparison of immunoglobulin G concentrations in primiparous and multiparous bovine colostrum.
      ), reductions in yield (1 kg) or increased colostrum dry matter (1%) were associated with marginal increases in IgG concentration (<3 g of IgG/L) (
      • Conneely M.
      • Berry D.P.
      • Sayers R.
      • Murphy J.P.
      • Lorenz I.
      • Doherty M.L.
      • Kennedy E.
      Factors associated with the concentration of immunoglobulin G in the colostrum of dairy cows.
      ,
      • Mann S.
      • Leal Yepes F.A.
      • Overton T.R.
      • Lock A.L.
      • Lamb S.V.
      • Wakshlag J.J.
      • Nydam D.V.
      Effect of dry period dietary energy level in dairy cattle on volume, concentrations of immunoglobulin G, insulin, and fatty acid composition of colostrum.
      ).

      Individual Variability

      It is known that colostrum yield and composition from dairy cows has high individual variation (
      • Kehoe S.I.
      • Jayarao B.M.
      • Heinrichs A.J.
      A survey of bovine colostrum composition and colostrum management practices on Pennsylvania dairy farms.
      ,
      • Baumrucker C.R.
      • Bruckmaier R.M.
      Colostrogenesis: IgG1 transcytosis mechanisms.
      ,
      • Dunn A.
      • Ashfield A.
      • Earley B.
      • Welsh M.
      • Gordon A.
      • Morrison S.J.
      Evaluation of factors associated with immunoglobulin G, fat, protein, and lactose concentrations in bovine colostrum and colostrum management practices in grassland-based dairy systems in Northern Ireland.
      ,
      • Borelli E.
      • Jonsson N.N.
      • Denholm K.S.
      Associations between putative risk factors and poor colostrum yield in Holstein Friesian cattle.
      ). Colostrum yield averaged (range) 4.3 (0 to 26.5) kg from Jersey (
      • Gavin K.
      • Neibergs H.
      • Hoffman A.
      • Kiser J.N.
      • Cornmesser M.A.
      • Haredasht S.A.
      • Martinez-Lopez B.
      • Wenz J.R.
      • Moore D.A.
      Low colostrum yield in Jersey cattle and potential risk factors.
      ) and 4.0–6.1 (0.0 to 43.8) kg from Holstein cows (
      • Borchardt S.
      • Sutter F.
      • Heuwieser W.
      • Venjakob P.
      Management-related factors in dry cows and their associations with colostrum quantity and quality on a large commercial dairy farm.
      ,
      • Rossi R.M.
      • Cullens F.M.
      • Bacigalupo P.
      • Sordillo L.M.
      • Abuelo A.
      Changes in biomarkers of metabolic stress during late gestation of dairy cows associated with colostrum volume and immunoglobulin content.
      ,
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      ), respectively with an intra-herd coefficient of variation of 25 to 74% (
      • Soufleri A.
      • Banos G.
      • Panousis N.
      • Fletouris D.
      • Arsenos G.
      • Kougioumtzis A.
      • Valergakis G.E.
      Evaluation of Factors Affecting Colostrum Quality and Quantity in Holstein Dairy Cattle.
      ). Yet, when considering the amount of colostrum needed to feed a calf 2 colostrum meals (for example 3–4 L at first feeding; 2 L at second feeding), 60.0 and 65.3% of Holstein cows failed to produce ≥ 6 L of first-milking colostrum (
      • Rossi R.M.
      • Cullens F.M.
      • Bacigalupo P.
      • Sordillo L.M.
      • Abuelo A.
      Changes in biomarkers of metabolic stress during late gestation of dairy cows associated with colostrum volume and immunoglobulin content.
      ,
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      ). Similarly, colostrum Brix % and IgG concentrations averaged (range) 23.8–27.6 (7.0 to 58.0) % (
      • Bielmann V.
      • Gillan J.
      • Perkins N.R.
      • Skidmore A.L.
      • Godden S.
      • Leslie K.E.
      An evaluation of Brix refractometry instruments for measurement of colostrum quality in dairy cattle.
      ,
      • Quigley J.D.
      • Lago A.
      • Chapman C.
      • Erickson P.
      • Polo J.
      Evaluation of the Brix refractometer to estimate immunoglobulin G concentration in bovine colostrum.
      ,
      • Gavin K.
      • Neibergs H.
      • Hoffman A.
      • Kiser J.N.
      • Cornmesser M.A.
      • Haredasht S.A.
      • Martinez-Lopez B.
      • Wenz J.R.
      • Moore D.A.
      Low colostrum yield in Jersey cattle and potential risk factors.
      ,
      • Borchardt S.
      • Sutter F.
      • Heuwieser W.
      • Venjakob P.
      Management-related factors in dry cows and their associations with colostrum quantity and quality on a large commercial dairy farm.
      ,
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      ) and 45.0–118.7 (1.4 to 261.2) g/L (
      • Gulliksen S.M.
      • Lie K.I.
      • Solverod L.
      • Osteras O.
      Risk factors associated with colostrum quality in Norwegian dairy cows.
      ,
      • Quigley J.D.
      • Lago A.
      • Chapman C.
      • Erickson P.
      • Polo J.
      Evaluation of the Brix refractometer to estimate immunoglobulin G concentration in bovine colostrum.
      ,
      • Dunn A.
      • Ashfield A.
      • Earley B.
      • Welsh M.
      • Gordon A.
      • Morrison S.J.
      Evaluation of factors associated with immunoglobulin G, fat, protein, and lactose concentrations in bovine colostrum and colostrum management practices in grassland-based dairy systems in Northern Ireland.
      ,
      • Shivley C.B.
      • Lombard J.E.
      • Urie N.J.
      • Haines D.M.
      • Sargent R.
      • Kopral C.A.
      • Earleywine T.J.
      • Olson J.D.
      • Garry F.B.
      Preweaned heifer management on US dairy operations: Part II. Factors associated with colostrum quality and passive transfer status of dairy heifer calves.
      ,
      • Kessler E.C.
      • Bruckmaier R.M.
      • Gross J.J.
      Colostrum composition and immunoglobulin G content in dairy and dual-purpose cattle breeds.
      ,
      • Rossi R.M.
      • Cullens F.M.
      • Bacigalupo P.
      • Sordillo L.M.
      • Abuelo A.
      Changes in biomarkers of metabolic stress during late gestation of dairy cows associated with colostrum volume and immunoglobulin content.
      ), respectively. Using the current industry standard for high-quality colostrum (IgG ≥ 50 g/L; Brix % ≥ 22.0%),
      • Buczinski S.
      • Vandeweerd J.
      Diagnostic accuracy of refractometry for assessing bovine colostrum quality: A systematic review and meta-analysis.
      and
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      reported that 7.7 to 32.7 and 21.5%, respectively of cows produced poor-quality colostrum. In addition, important nutritional components for the calf including colostral fat [mean (SD) [Quartile 1, Quartile 3]; 6.4 (3.3) [2.5, 10.9] %], protein [17.8 (4.0) [12.7, 22.7] %], and lactose [2.15 (0.73) [1.2, 3.1] %] vary by cow (
      • Soufleri A.
      • Banos G.
      • Panousis N.
      • Fletouris D.
      • Arsenos G.
      • Kougioumtzis A.
      • Valergakis G.E.
      Evaluation of Factors Affecting Colostrum Quality and Quantity in Holstein Dairy Cattle.
      ) resulting in an inconsistent nutritional value.
      Researchers investigating factors that affect colostrum yield and quality have mostly used observational data to uncover individual animal factors such as parity (
      • Conneely M.
      • Berry D.P.
      • Sayers R.
      • Murphy J.P.
      • Lorenz I.
      • Doherty M.L.
      • Kennedy E.
      Factors associated with the concentration of immunoglobulin G in the colostrum of dairy cows.
      ,
      • Dunn A.
      • Ashfield A.
      • Earley B.
      • Welsh M.
      • Gordon A.
      • Morrison S.J.
      Evaluation of factors associated with immunoglobulin G, fat, protein, and lactose concentrations in bovine colostrum and colostrum management practices in grassland-based dairy systems in Northern Ireland.
      ,
      • Fischer-Tlustos A.
      • Hertogs K.
      • van Niekerk J.
      • Nagorske M.
      • Haines D.
      • Steele M.
      Oligosaccharide concentrations in colostrum, transition milk, and mature milk of primi-and multiparous Holstein cows during the first week of lactation.
      ,
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      ), breed (
      • Muller L.
      • Ellinger D.
      Colostral immunoglobulin concentrations among breeds of dairy cattle.
      ,
      • Guy M.A.
      • McFadden T.B.
      • Cockrell D.C.
      • Besser T.E.
      Regulation of colostrum formation in beef and dairy cows.
      ,
      • Zarcula S.
      • Cernescu H.
      • Mircu C.
      • Tulcan C.
      • Morvay A.
      • Baul S.
      • Popovici D.
      Influence of breed, parity and food intake on chemical composition of first colostrum in cow.
      ,
      • Kessler E.C.
      • Bruckmaier R.M.
      • Gross J.J.
      Colostrum composition and immunoglobulin G content in dairy and dual-purpose cattle breeds.
      ), and month of calving (
      • Dunn A.
      • Ashfield A.
      • Earley B.
      • Welsh M.
      • Gordon A.
      • Morrison S.J.
      Evaluation of factors associated with immunoglobulin G, fat, protein, and lactose concentrations in bovine colostrum and colostrum management practices in grassland-based dairy systems in Northern Ireland.
      ,
      • Zarei S.
      • Ghorbani G.R.
      • Khorvash M.
      • Martin O.B.
      • Mahdavi A.H.
      • Riasi A.
      The impact of season, parity, and volume of colostrum on Holstein dairy cows colostrum composition.
      ,
      • Soufleri A.
      • Banos G.
      • Panousis N.
      • Fletouris D.
      • Arsenos G.
      • Kougioumtzis A.
      • Valergakis G.E.
      Evaluation of Factors Affecting Colostrum Quality and Quantity in Holstein Dairy Cattle.
      ,
      • Rossi R.M.
      • Cullens F.M.
      • Bacigalupo P.
      • Sordillo L.M.
      • Abuelo A.
      Changes in biomarkers of metabolic stress during late gestation of dairy cows associated with colostrum volume and immunoglobulin content.
      ) that commonly arise as associated with colostrum production. Within common dairy breeds, Jersey cows produced colostrum with the highest quality but experience periods of low colostrum supply specifically during the fall and winter months (
      • Muller L.
      • Ellinger D.
      Colostral immunoglobulin concentrations among breeds of dairy cattle.
      ,
      • Gavin K.
      • Neibergs H.
      • Hoffman A.
      • Kiser J.N.
      • Cornmesser M.A.
      • Haredasht S.A.
      • Martinez-Lopez B.
      • Wenz J.R.
      • Moore D.A.
      Low colostrum yield in Jersey cattle and potential risk factors.
      ). Multiparous cows produced a greater volume of colostrum with higher IgG as well as protein concentrations while fat concentration was lower compared with primiparous cows (
      • Karl M.
      • Staufenbiel R.
      Factors influencing the amount of initial colostrum in Holstein-Friesian cows and their relationship to postpartum calcium concentration.
      ,
      • Dunn A.
      • Ashfield A.
      • Earley B.
      • Welsh M.
      • Gordon A.
      • Morrison S.J.
      Evaluation of factors associated with immunoglobulin G, fat, protein, and lactose concentrations in bovine colostrum and colostrum management practices in grassland-based dairy systems in Northern Ireland.
      ,
      • Soufleri A.
      • Banos G.
      • Panousis N.
      • Fletouris D.
      • Arsenos G.
      • Kougioumtzis A.
      • Valergakis G.E.
      Evaluation of Factors Affecting Colostrum Quality and Quantity in Holstein Dairy Cattle.
      ,
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      ). Other variables from selected studies such as characteristics of the previous lactation or calf as well as the heritability of colostrum production are summarized in Table 1. Recently, authors have associated carrying a heifer calf as well as having a stillbirth with a lower colostrum yield in both Holstein and Jersey cattle (
      • Karl M.
      • Staufenbiel R.
      Factors influencing the amount of initial colostrum in Holstein-Friesian cows and their relationship to postpartum calcium concentration.
      ,
      • Gavin K.
      • Neibergs H.
      • Hoffman A.
      • Kiser J.N.
      • Cornmesser M.A.
      • Haredasht S.A.
      • Martinez-Lopez B.
      • Wenz J.R.
      • Moore D.A.
      Low colostrum yield in Jersey cattle and potential risk factors.
      ,
      • Borchardt S.
      • Sutter F.
      • Heuwieser W.
      • Venjakob P.
      Management-related factors in dry cows and their associations with colostrum quantity and quality on a large commercial dairy farm.
      ,
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      ) and these associations might be influenced by calf birth weight (
      • Conneely M.
      • Berry D.P.
      • Sayers R.
      • Murphy J.P.
      • Lorenz I.
      • Doherty M.L.
      • Kennedy E.
      Factors associated with the concentration of immunoglobulin G in the colostrum of dairy cows.
      ,
      • Sutter F.
      • Borchardt S.
      • Schuenemann G.M.
      • Rauch E.
      • Erhard M.
      • Heuwieser W.
      Evaluation of 2 different treatment procedures after calving to improve harvesting of high-quantity and high-quality colostrum.
      ). Circulating concentrations of placental lactogen during gestation has been positively associated with calf birth weight and milk production in the subsequent lactation (
      • Bolander Jr., F.
      • Ulberg L.
      • Fellows R.
      Circulating placental lactogen levels in dairy and beef cattle.
      ,
      • Hayden T.
      • Thomas C.
      • Forsyth I.A.
      Effect of number of young born (litter size) on milk yield of goats: role for placental lactogen.
      ,
      • Patel O.
      • Hirako M.
      • Takahashi T.
      • Sasaki N.
      • Domeki I.
      Plasma bovine placental lactogen concentration throughout pregnancy in the cow; relationship to stage of pregnancy, fetal mass, number and postpartum milk yield.
      ) and circulating progesterone and estrogen among other hormones have been reviewed for their effect on mammary development (
      • Erb R.
      Harmonal control of mammogenesis and onset of lactation in cows–A Review.
      ). We hypothesize that endocrine signals during late gestation might contribute to the association between calf related variables (sex, birth weight, and stillbirth) and colostrum yield.
      Table 1Previous lactation and calf characteristics as well as heritability associated with colostrum yield and quality
      ReferenceVariableYieldQuality
      Kessler et al., 2014305-d cumulative milk yieldNot associated
      Cabral et al., 2016Previous lactation milk yieldNot associated
      Dunn et al., 2017Previous lactation 305-d milk yield
      Kessler et al., 2020bPrevious lactation 305-d milk yieldNot associated
      Poindexter, 2021Milk yield for first 70 DIM
      Borchardt et al., 2022Previous lactation 305-d milk yield
      Westhoff et al., 2023bPrevious lactation 305ME, kg
      ≤13,090ReferentReferent
      13,091–15862ND
      >15,862
      Cabral et al., 2016Previous lactation lengthNot associated
      Gavin et al., 2018Previous lactation lengthNot associated
      Westhoff et al., 2023bPrevious lactation length, dNot associated
      ≤297Referent
      297–344nd
      >344
      Kessler et al., 2020bGestation lengthNot associated
      Borchardt et al., 2022Gestation length
      Short = 257–269, normal = 270–280, long = 281–293.
      Not associatedNot associated
      Short
      Normal
      Long
      Westhoff et al., 2023bGestation length
      Short = 263–273, normal = 274–282, long = 283–293.
      ShortReferentReferent
      NormalND
      LongND
      Angulo et al., 2015Calf sex
      FemaleReferentReferent
      Male
      Gavin et al., 2018Calf sex
      TwinsReferentReferent
      Female
      MaleND
      Kessler et al., 2020bCalf sexNot associatedNot associated
      Poindexter, 2021Calf sex
      FemaleReferent
      Male
      Twins
      Borchardt et al., 2022Calf sexNot associated
      FemaleReferent
      Male
      Twins
      Westhoff et al., 2023bCalf sex
      FemaleReferentReferent
      Male
      TwinsND
      Karl and Staufenbiel, 2016, 2017Stillbirth
      AliveReferentReferent
      Dead
      Borchardt et al., 2022StillbirthNot associated
      AliveReferent
      Dead
      Westhoff et al., 2023bStillbirth
      AliveReferentReferent
      Dead
      Conneely et al., 2013Calf birth weightNot associated
      Karl and Staufenbiel, 2016, 2017Calf birth weightNot associated
      Kessler et al., 2020bCalf birth weightNot associated
      Soufleri et al., 2019HeritabilityNot associated
      Costa et al., 2021bHeritability
      Arrows indicate a positive or negative association with the variable relative to the referent when variable was categorized. Not associated = P > 0.05, nd = no difference from referent group (P > 0.05), dash = not reported.
      1 Short = 257–269, normal = 270–280, long = 281–293.
      2 Short = 263–273, normal = 274–282, long = 283–293.
      Other variables such as gestation length, previous lactation length, and milk yield in the current or previous lactation resulted in mixed associations with colostrum production (
      • Kessler E.C.
      • Bruckmaier R.M.
      • Gross J.J.
      Milk production during the colostral period is not related to the later lactational performance in dairy cows.
      ,
      • Cabral R.G.
      • Chapman C.E.
      • Aragona K.M.
      • Clark E.
      • Lunak M.
      • Erickson P.S.
      Predicting colostrum quality from performance in the previous lactation and environmental changes.
      ,
      • Gavin K.
      • Neibergs H.
      • Hoffman A.
      • Kiser J.N.
      • Cornmesser M.A.
      • Haredasht S.A.
      • Martinez-Lopez B.
      • Wenz J.R.
      • Moore D.A.
      Low colostrum yield in Jersey cattle and potential risk factors.
      ,
      • Kessler E.C.
      • Pistol G.C.
      • Bruckmaier R.M.
      • Gross J.J.
      Pattern of milk yield and immunoglobulin concentration and factors associated with colostrum quality at the quarter level in dairy cows after parturition.
      ,
      • Poindexter M.
      Use of Vitamin D and choline to improve health and productivy of dairy cows and their offspring. in Department of Animal Science.
      ,
      • Borchardt S.
      • Sutter F.
      • Heuwieser W.
      • Venjakob P.
      Management-related factors in dry cows and their associations with colostrum quantity and quality on a large commercial dairy farm.
      ,
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      ) and might be the result of differences in data collection, statistical analysis, or indicate the lack of a cause-effect relationship. Colostrum quality and composition appear to have low to moderate heritability (
      • Soufleri A.
      • Banos G.
      • Panousis N.
      • Fletouris D.
      • Arsenos G.
      • Valergakis G.
      Genetic parameters of colostrum traits in Holstein dairy cows.
      ,
      • Costa A.
      • Franzoi M.
      • Visentin G.
      • Goi A.
      • De Marchi M.
      • Penasa M.
      The concentrations of immunoglobulins in bovine colostrum determined by the gold standard method are genetically correlated with their near-infrared prediction.
      ) and low colostrum yield has been linked to Holstein and Jersey sire lines (
      • Karl M.
      • Staufenbiel R.
      Factors influencing the amount of initial colostrum in Holstein-Friesian cows and their relationship to postpartum calcium concentration.
      ,
      • Gavin K.
      • Neibergs H.
      • Hoffman A.
      • Kiser J.N.
      • Cornmesser M.A.
      • Haredasht S.A.
      • Martinez-Lopez B.
      • Wenz J.R.
      • Moore D.A.
      Low colostrum yield in Jersey cattle and potential risk factors.
      ). However, the feasibility and extent to which genetic selection might improve colostrum production warrants further investigation.
      An emerging area of study has considered the effect of the dam's metabolism on colostrum production. In an observational study,
      • Poindexter M.
      Use of Vitamin D and choline to improve health and productivy of dairy cows and their offspring. in Department of Animal Science.
      observed a positive association between colostrum yield and hypocalcemia when circulating calcium was measured at 1 DIM. Further, it was recently revealed that production of ≥6 L of colostrum was associated with an elevated prepartum β-hydroxybutyrate concentration and antioxidant potential as well as a lower cholesterol concentration and oxidant status index in Holstein dairy cows (
      • Rossi R.M.
      • Cullens F.M.
      • Bacigalupo P.
      • Sordillo L.M.
      • Abuelo A.
      Changes in biomarkers of metabolic stress during late gestation of dairy cows associated with colostrum volume and immunoglobulin content.
      ). Elevated colostral IgG concentration or Brix % was associated with higher prepartum serum albumin and glucose concentrations as well as a lower calcium concentration and glutamate dehydrogenase activity, as well as urinary net acid base excretion (
      • Immler M.
      • Failing K.
      • Gärtner T.
      • Wehrend A.
      • Donat K.
      Associations between the metabolic status of the cow and colostrum quality as determined by Brix refractometry.
      ,
      • Rossi R.M.
      • Cullens F.M.
      • Bacigalupo P.
      • Sordillo L.M.
      • Abuelo A.
      Changes in biomarkers of metabolic stress during late gestation of dairy cows associated with colostrum volume and immunoglobulin content.
      ). Concentrations of circulating fatty acids as well as β-hydroxybutyrate at 0, 1, and 7 DIM were positively associated with colostrum yield (
      • Karl M.
      • Staufenbiel R.
      Factors influencing the immunoglobulin concentration in the first colostrum in Holstein-Friesian dairy cows and their relationship to the postpartum calcium concentration in blood and colostrum.
      ) and for every one liter increase in colostrum yield above the mean of 5.43 L, cows had a 1.1 (1.0–1.1) greater odds of hyperketonemia when evaluated between 7 and 14 DIM (
      • Vanholder T.
      • Papen J.
      • Bemers R.
      • Vertenten G.
      • Berge A.
      Risk factors for subclinical and clinical ketosis and association with production parameters in dairy cows in the Netherlands.
      ). Further, a 10.1–15.0% prevalence of hyperketonemia, at the herd level, during the early postpartum period (3–14 DIM) was associated with a greater colostrum yield (
      • Westhoff T.
      • Overton T.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: Prepartum nutrition and metabolic indicators.
      ). While these data are suggestive of a relationship between maternal metabolism and colostrum production in Holstein dairy cattle, the causality as well as the interactions between management, nutrition, and metabolism in respect to colostrogenesis remain unknown.

      Herd-Level Variability

      Herd-level differences in colostrum yield and composition within a geographic region (
      • Costa A.
      • Goi A.
      • Penasa M.
      • Nardino G.
      • Posenato L.
      • De Marchi M.
      Variation of immunoglobulins G, A, and M and bovine serum albumin concentration in Holstein cow colostrum.
      ,
      • Rossi R.M.
      • Cullens F.M.
      • Bacigalupo P.
      • Sordillo L.M.
      • Abuelo A.
      Changes in biomarkers of metabolic stress during late gestation of dairy cows associated with colostrum volume and immunoglobulin content.
      ,
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      ) suggest variables beyond season of calving and individual animal factors influence colostrum synthesis. Annual median (Q1, Q3) colostrum yield and mean (SD) Brix % ranged from 3.7 (2.6, 5.3) to 7.7 (6.0, 8.6) kg and 22.0 (2.7) to 28.4 (5.1) % on 18 Holstein dairy farms (
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      ). Although less than the intra-herd variability, inter-herd coefficient of variation was 35% for colostrum yield and between 6 and 22% for colostrum components (
      • Soufleri A.
      • Banos G.
      • Panousis N.
      • Fletouris D.
      • Arsenos G.
      • Kougioumtzis A.
      • Valergakis G.E.
      Evaluation of Factors Affecting Colostrum Quality and Quantity in Holstein Dairy Cattle.
      ) suggesting farm management, prepartum nutrition, or environmental conditions influence colostrum synthesis. Recent insights into the influence of dry period management (
      • Grusenmeyer D.
      • Ryan C.
      • Galton D.
      • Overton T.
      Shortening the dry period from 60 to 40 days does not affect colostrum quality but decreases colostrum yield by Holstein cows.
      ,
      • Mayasari N.
      • de Vries Reilingh G.
      • Nieuwland M.G.
      • Remmelink G.J.
      • Parmentier H.K.
      • Kemp B.
      • van Knegsel A.T.
      Effect of maternal dry period length on colostrum immunoglobulin content and on natural and specific antibody titers in calves.
      ,
      • Borchardt S.
      • Sutter F.
      • Heuwieser W.
      • Venjakob P.
      Management-related factors in dry cows and their associations with colostrum quantity and quality on a large commercial dairy farm.
      ,
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      ) and prepartum nutrition (
      • Martinez N.
      • Rodney R.
      • Block E.
      • Hernandez L.
      • Nelson C.
      • Lean I.
      • Santos J.
      Effects of prepartum dietary cation-anion difference and source of vitamin D in dairy cows: Lactation performance and energy metabolism.
      ,
      • Swartz T.H.
      • Bradford B.J.
      • Malysheva O.
      • Caudill M.A.
      • Mamedova L.K.
      • Estes K.A.
      Effects of dietary rumen-protected choline supplementation on colostrum yields, quality, and choline metabolites from dairy cattle.
      ,
      • Hare K.S.
      • Fischer-Tlustos A.J.
      • Wood K.M.
      • Cant J.P.
      • Steele M.A.
      Prepartum nutrient intake and colostrum yield and composition in ruminants.
      ,
      • Westhoff T.
      • Overton T.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: Prepartum nutrition and metabolic indicators.
      ), emphasize the potential to alter colostrum synthesis. However, because of the individuality and seasonality of colostrum production as well as the lack of complete knowledge on the metabolic and endocrine signals that regulate colostrum synthesis (
      • Baumrucker C.R.
      • Macrina A.L.
      • Bruckmaier R.M.
      Colostrogenesis: Role and Mechanism of the Bovine Fc Receptor of the Neonate (FcRn).
      ,
      • Baumrucker C.R.
      • Gross J.J.
      • Bruckmaier R.M.
      The importance of colostrum in maternal care and its formation in mammalian species.
      ), a lack of understanding of mechanisms underlying colostrogenesis have largely limited our ability to explain the aforementioned differences in colostrum yield or composition. As such, our success in the ability to employ on-farm management and nutritional strategies to improve colostrum yield and composition has been minimal thus far and highlights the need for mechanistic studies to fill this important knowledge gap.

      PREPARTUM NUTRITIONAL AND MANAGEMENT STRATEGIES

      Prepartum Nutrition

      Prepartum nutritional strategies are often evaluated by their effect on postpartum health and productivity. However, with the growing interest in and utility for colostrum (
      • Lopez A.J.
      • Heinrichs A.J.
      Invited review: The importance of colostrum in the newborn dairy calf.
      ,
      • Costa A.
      • Sneddon N.
      • Goi A.
      • Visentin G.
      • Mammi L.
      • Savarino E.
      • Zingone F.
      • Formigoni A.
      • Penasa M.
      • De Marchi M.
      Invited review: Bovine colostrum, a promising ingredient for humans and animals—Properties, processing technologies, and uses.
      ), researchers have recently started to simultaneously evaluate the effect of these nutritional strategies on colostrum production. We encourage consideration of colostrum outcomes in these types of studies as future meta-analyses can be conducted to provide clarification of effect sizes and directions where there is currently no consensus.

      Dietary energy

      For a comprehensive review of the current knowledge on the effect of prepartum intake of carbohydrates, fat, and protein on colostrum production, we direct the reader to
      • Hare K.S.
      • Fischer-Tlustos A.J.
      • Wood K.M.
      • Cant J.P.
      • Steele M.A.
      Prepartum nutrient intake and colostrum yield and composition in ruminants.
      . In brief, altering starch concentrations to increase the energy density of the prepartum diets does not appear to alter colostrum yield (
      • Mann S.
      • Leal Yepes F.A.
      • Overton T.R.
      • Lock A.L.
      • Lamb S.V.
      • Wakshlag J.J.
      • Nydam D.V.
      Effect of dry period dietary energy level in dairy cattle on volume, concentrations of immunoglobulin G, insulin, and fatty acid composition of colostrum.
      ,
      • Richards B.
      • Janovick N.
      • Moyes K.
      • Beever D.
      • Drackley J.
      Comparison of prepartum low-energy or high-energy diets with a 2-diet far-off and close-up strategy for multiparous and primiparous cows.
      ,
      • Vasquez J.
      • McCarthy M.
      • Richards B.
      • Perfield K.
      • Carlson D.
      • Lock A.
      • Drackley J.
      Effects of prepartum diets varying in dietary energy density and monensin on early-lactation performance in dairy cows.
      ) but resulted in a greater colostrum IgG and insulin concentrations as well as altered fatty acid composition (
      • Mann S.
      • Leal Yepes F.A.
      • Overton T.R.
      • Lock A.L.
      • Lamb S.V.
      • Wakshlag J.J.
      • Nydam D.V.
      Effect of dry period dietary energy level in dairy cattle on volume, concentrations of immunoglobulin G, insulin, and fatty acid composition of colostrum.
      ,
      • Fischer-Tlustos A.
      • Hare K.
      • Haisan J.
      • Shi W.
      • Cant J.
      • Oba M.
      • Steele M.
      Transition diet starch content impacts colostrum and transition milk composition and immunoglobulin G and insulin concentrations in Holstein dairy cattle.
      ). Inclusion of fat in the prepartum diet did not affect yield or component concentrations of colostrum and the effect of fat supplementation on IgG concentration remains mixed (
      • Garcia M.
      • Greco L.F.
      • Favoreto M.G.
      • Marsola R.S.
      • Martins L.T.
      • Bisinotto R.S.
      • Shin J.H.
      • Lock A.L.
      • Block E.
      • Thatcher W.W.
      • Santos J.E.
      • Staples C.R.
      Effect of supplementing fat to pregnant nonlactating cows on colostral fatty acid profile and passive immunity of the newborn calf.
      ,
      • Salehi R.
      • Ambrose D.J.
      • Oba M.
      Short communication: Effects of prepartum diets supplemented with rolled oilseeds on Brix values and fatty acid profile of colostrum.
      ,
      • Jolazadeh A.R.
      • Mohammadabadi T.
      • Dehghan-banadaky M.
      • Chaji M.
      • Garcia M.
      Effect of supplementing calcium salts of n-3 and n-6 fatty acid to pregnant nonlactating cows on colostrum composition, milk yield, and reproductive performance of dairy cows.
      ,
      • Daneshvar D.
      • Hashemzadeh F.
      • Ghasemi E.
      • Khorvash M.
      Effects of stearic acid supplementation and starch concentration in close-up diets on performance and metabolic responses of transition dairy cows.
      ,
      • Sun X.
      • Hou Y.
      • Wang Y.
      • Guo C.
      • Wang Q.
      • Zhang Y.
      • Yang Z.
      • Wang Z.
      • Cao Z.
      • Wang W.
      • Li S.
      The Blood Immune Cell Count, Immunoglobulin, Inflammatory Factor, and Milk Trace Element in Transition Cows and Calves Were Altered by Increasing the Dietary n-3 or n-6 Polyunsaturated Fatty Acid Levels.
      ,
      • Sun X.
      • Guo C.
      • Zhang Y.
      • Wang Q.
      • Yang Z.
      • Wang Z.
      • Wang W.
      • Cao Z.
      • Niu M.
      • Li S.
      Effect of diets enriched in n-6 or n-3 fatty acids on dry matter intake, energy balance, oxidative stress, and milk fat profile of transition cows.
      ).

      Dietary protein

      Colostrum yield from multiparous cows did not differ when shifting the prepartum metabolizable protein (MP) supply from 744 to 976 or 849 to 1,200 to 1,387 g of estimated MP/d (
      • Farahani T.A.
      • Amanlou H.
      • Kazemi-Bonchenari M.
      Effects of shortening the close-up period length coupled with increased supply of metabolizable protein on performance and metabolic status of multiparous Holstein cows.
      ,
      • Farahani T.A.
      • Amanlou H.
      • Farsuni N.
      • Kazemi-Bonchenari M.
      Interactions of protein levels fed to Holstein cows pre-and postpartum on productive and metabolic responses.
      ). In a study with 2 levels of MP (65 vs. 90 g/kg DM) and 2 levels of DMI,
      • Akhtar M.
      • Ahmed M.
      • Pasha T.
      • Abdullah M.
      • Shahbakht R.
      • Haq M.
      • Haque M.
      Colostrum production, calf birth weight, and postpartum ovarian follicular activity of dairy cows fed restricted diet with different protein levels during the prepartum period.
      did not observe an effect of MP level, DMI, or the interaction on colostrum yield, IgG, or component concentrations. Similarly, the level of crude protein fed prepartum (
      • Santos J.
      • DePeters E.
      • Jardon P.
      • Huber J.
      Effect of prepartum dietary protein level on performance of primigravid and multiparous Holstein dairy cows.
      ,
      • Toghyani E.
      • Moharrery A.
      Effect of Various Levels of Dietary Protein in Transition Period on Colostrum Quality and Serum Immunoglobulin Concentration in Holstein Cows and their Newborn Calves.
      ) or inclusion of rumen-protected lysine (
      • Fehlberg L.
      • Guadagnin A.
      • Thomas B.
      • Sugimoto Y.
      • Shinzato I.
      • Cardoso F.
      Feeding rumen-protected lysine prepartum increases energy-corrected milk and milk component yields in Holstein cows during early lactation.
      ) did not affect colostrum composition. However, we recently observed a tendency for an interaction between MP level fed prepartum and parity group such that cows entering parity 2 tended to produce more colostrum (9.4 ± 0.9 vs. 7.2 ± 0.9 kg) when fed an elevated level of MP (1,606 vs. 1,180 g of estimated MP/d) but colostrum yield from parity ≥ 3 cows was not affected by MP supply (5.1 ± 1.0 vs. 6.4 ± 1.0 kg; Westhoff et al., personal communication), respectively. Further, increasing the MP supply during the far-off (1,203 vs. 846 g of estimated MP/d) and close-up (1,631 vs. 1,258 g of estimated MP/d) periods resulted in a treatment x parity interaction for IgG concentration such that cows entering parity 2 fed an elevated MP supply had a greater IgG concentration (61.3 ± 2.3 vs. 55.2 ± 2.8 g/L) compared with parity 2 cows fed a lower MP supply, but MP supply did not affect IgG concentration in parity ≥ 3 cows (58.4 ± 3.0 vs. 56.8 ± 2.9 g/L;
      • Van Hese I.
      • Goossens K.
      • Vandaele L.
      • Ampe B.
      • Haegeman A.
      • Opsomer G.
      The effect of maternal supply of rumen-protected protein to Holstein Friesian cows during the dry period on the transfer of passive immunity and colostral microbial composition.
      ), respectively. Although nulliparous heifers were not included in the 2 aforementioned studies, it remains plausible that younger cows might benefit from an additional MP supply to support mammary epithelial cell turnover during the dry period; however, the mechanism responsible as well as the validity of the treatment x parity interaction remain uncertain. Because of limited data as well as the aforementioned interaction, the effect of prepartum MP supply on colostrum production should be further investigated with inclusion of nulliparous heifers.

      Hypocalcemia prevention strategies

      Prepartum dietary strategies to mitigate hypocalcemia, including feeding a zeolite or manipulating the dietary calcium or cation anion difference (DCAD), alter prepartum DMI and circulating mineral concentrations (
      • Kerwin A.L.
      • Ryan C.M.
      • Leno B.M.
      • Jakobsen M.
      • Theilgaard P.
      • Barbano D.M.
      • Overton T.R.
      Effects of feeding synthetic zeolite A during the prepartum period on serum mineral concentration, oxidant status, and performance of multiparous Holstein cows.
      ,
      • Santos J.E.P.
      • Lean I.J.
      • Golder H.
      • Block E.
      Meta-analysis of the effects of prepartum dietary cation-anion difference on performance and health of dairy cows.
      ). Because of the metabolic demand of colostrum synthesis (
      • Goff J.P.
      • Horst R.L.
      Physiological Changes at Parturition and Their Relationship to Metabolic Disorders.
      ), disruptions in prepartum mineral metabolism or nutrient intake have potential to interfere with colostrogenesis. When feeding a diet including a zeolite at 500 g/d, colostrum yield (5.8 ± 0.8 vs. 7.3 ± 0.8 kg) and IgG concentrations (83.4 ± 4.8 vs. 76.3 ± 4.8 g/L) did not differ in the treatment compared with the control diet, respectively (
      • Kerwin A.L.
      • Ryan C.M.
      • Leno B.M.
      • Jakobsen M.
      • Theilgaard P.
      • Barbano D.M.
      • Overton T.R.
      Effects of feeding synthetic zeolite A during the prepartum period on serum mineral concentration, oxidant status, and performance of multiparous Holstein cows.
      ). Further, colostrum fat, protein, and lactose concentrations were not affected by a zeolite fed at 150 or 300 g/d (
      • Marin M.P.
      • Pogurschi E.N.
      • Marin I.
      • Nicolae C.G.
      Influence of natural zeolites supplemented with inorganic selenium on the productive performance of dairy cows.
      ). Apart from an elevated IgG concentration from cows fed a negative (−22 mEq/100 g) compared with a neutral (−3 mEq/100 g) DCAD diet described by
      • Diehl A.
      • Bernard J.
      • Tao S.
      • Smith T.
      • Marins T.
      • Kirk D.
      • McLean D.
      • Chapman J.
      Blood mineral and gas concentrations of calves born to cows fed prepartum diets differing in dietary cation-anion difference and calcium concentration.
      , altering the prepartum DCAD (
      • Lopera C.
      • Zimpel R.
      • Vieira-Neto A.
      • Lopes F.R.
      • Ortiz W.
      • Poindexter M.
      • Faria B.N.
      • Gambarini M.L.
      • Block E.
      • Nelson C.D.
      • Santos J.E.P.
      Effects of level of dietary cation-anion difference and duration of prepartum feeding on performance and metabolism of dairy cows.
      ,
      • Martinez N.
      • Rodney R.
      • Block E.
      • Hernandez L.
      • Nelson C.
      • Lean I.
      • Santos J.
      Effects of prepartum dietary cation-anion difference and source of vitamin D in dairy cows: Lactation performance and energy metabolism.
      ,
      • Zimpel R.
      • Almeida K.V.
      • Marinho M.N.
      • Ruiz A.R.
      • Perdomo M.C.
      • Poindexter M.B.
      • Husnain A.
      • Vieira-Neto A.
      • Silva A.C.M.
      • Arshad U.
      • Nelson C.D.
      • Santos J.E.P.
      Prepartum level of dietary cation-anion difference fed to nulliparous cows: Lactation and reproductive responses.
      ) or prepartum DCAD and dietary calcium concentration (
      • Glosson K.M.
      • Zhang X.
      • Bascom S.S.
      • Rowson A.D.
      • Wang Z.
      • Drackley J.K.
      Negative dietary cation-anion difference and amount of calcium in prepartum diets: Effects on milk production, blood calcium, and health.
      ,
      • Rajaeerad A.
      • Ghorbani G.R.
      • Khorvash M.
      • Sadeghi-Sefidmazgi A.
      • Mahdavi A.H.
      • Rashidi S.
      • Wilkens M.R.
      • Hünerberg M.
      Impact of a ration negative in dietary cation–anion difference and varying calcium supply fed before calving on colostrum quality of the dams and health status and growth performance of the calves.
      ,
      • Graef G.M.
      • Kerwin A.L.
      • Ferro L.N.
      • Ordaz-Puga S.
      • Ryan C.M.
      • Westhoff T.A.
      • Barbano D.M.
      • Glosson K.M.
      • Zanzalari K.P.
      • Chapman J.D.
      • Overton T.R.
      Effect of Prepartum DCAD Strategy and level of Dietary Calcium on Postpartum Calcium Status and Performacne of Multiparous Holstein Cows.
      ) did not affect colostrum IgG concentration or Brix %. While most authors did not observe an effect on colostrum yield (
      • Weich W.
      • Block E.
      • Litherland N.
      Extended negative dietary cation-anion difference feeding does not negatively affect postpartum performance of multiparous dairy cows.
      ,
      • Diehl A.
      • Bernard J.
      • Tao S.
      • Smith T.
      • Marins T.
      • Kirk D.
      • McLean D.
      • Chapman J.
      Blood mineral and gas concentrations of calves born to cows fed prepartum diets differing in dietary cation-anion difference and calcium concentration.
      ,
      • Martinez N.
      • Rodney R.
      • Block E.
      • Hernandez L.
      • Nelson C.
      • Lean I.
      • Santos J.
      Effects of prepartum dietary cation-anion difference and source of vitamin D in dairy cows: Lactation performance and energy metabolism.
      ,
      • Glosson K.M.
      • Zhang X.
      • Bascom S.S.
      • Rowson A.D.
      • Wang Z.
      • Drackley J.K.
      Negative dietary cation-anion difference and amount of calcium in prepartum diets: Effects on milk production, blood calcium, and health.
      ,
      • Rajaeerad A.
      • Ghorbani G.R.
      • Khorvash M.
      • Sadeghi-Sefidmazgi A.
      • Mahdavi A.H.
      • Rashidi S.
      • Wilkens M.R.
      • Hünerberg M.
      Impact of a ration negative in dietary cation–anion difference and varying calcium supply fed before calving on colostrum quality of the dams and health status and growth performance of the calves.
      ,
      • Zimpel R.
      • Almeida K.V.
      • Marinho M.N.
      • Ruiz A.R.
      • Perdomo M.C.
      • Poindexter M.B.
      • Husnain A.
      • Vieira-Neto A.
      • Silva A.C.M.
      • Arshad U.
      • Nelson C.D.
      • Santos J.E.P.
      Prepartum level of dietary cation-anion difference fed to nulliparous cows: Lactation and reproductive responses.
      ), feeding diets with a more severe negative DCAD has been associated with reduced (
      • Lopera C.
      • Zimpel R.
      • Vieira-Neto A.
      • Lopes F.R.
      • Ortiz W.
      • Poindexter M.
      • Faria B.N.
      • Gambarini M.L.
      • Block E.
      • Nelson C.D.
      • Santos J.E.P.
      Effects of level of dietary cation-anion difference and duration of prepartum feeding on performance and metabolism of dairy cows.
      ,
      • Westhoff T.
      • Overton T.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: Prepartum nutrition and metabolic indicators.
      ) and increased colostrum yield (
      • Graef G.M.
      • Kerwin A.L.
      • Ferro L.N.
      • Ordaz-Puga S.
      • Ryan C.M.
      • Westhoff T.A.
      • Barbano D.M.
      • Glosson K.M.
      • Zanzalari K.P.
      • Chapman J.D.
      • Overton T.R.
      Effect of Prepartum DCAD Strategy and level of Dietary Calcium on Postpartum Calcium Status and Performacne of Multiparous Holstein Cows.
      ). Although the reduction in yield was hypothesized to be affected by a lower DMI (
      • Lopera C.
      • Zimpel R.
      • Vieira-Neto A.
      • Lopes F.R.
      • Ortiz W.
      • Poindexter M.
      • Faria B.N.
      • Gambarini M.L.
      • Block E.
      • Nelson C.D.
      • Santos J.E.P.
      Effects of level of dietary cation-anion difference and duration of prepartum feeding on performance and metabolism of dairy cows.
      ), the complexity of calcium homeostasis, including the effect of 25-hydroxyvitamin D3 (discussed below), the role of the different widely used dietary hypocalcemia prevention strategies, and serotonin's negative association with colostrum yield (
      • Hernández-Castellano L.E.
      • Hernandez L.L.
      • Weaver S.
      • Bruckmaier R.M.
      Increased serum serotonin improves parturient calcium homeostasis in dairy cows.
      ,
      • Kessler E.C.
      • Wall S.K.
      • Hernandez L.L.
      • Bruckmaier R.M.
      • Gross J.J.
      Short communication: Circulating serotonin is related to the metabolic status and lactational performance at the onset of lactation in dairy cows.
      ), deserve further attention.

      Mineral and vitamin inclusion

      Recent evidence suggests the source of dietary vitamin D influences colostrum synthesis.
      • Martinez N.
      • Rodney R.
      • Block E.
      • Hernandez L.
      • Nelson C.
      • Lean I.
      • Santos J.
      Effects of prepartum dietary cation-anion difference and source of vitamin D in dairy cows: Lactation performance and energy metabolism.
      observed that feeding 3 mg of calcidiol (25-hydroxyvitamin D3) tended to increase colostrum yield (7.8 ± 0.8 vs. 6.0 ± 0.8 kg) compared with cows fed cholecalciferol (vitamin D3), respectively. Further, when fed in combination with a positive DCAD diet, calcidiol also tended to increase concentrations of fat, protein, and total solids; however, feeding calcidiol did not result in altered colostral fat, protein, or total solids concentrations when cows were fed a diet with a negative DCAD (
      • Martinez N.
      • Rodney R.
      • Block E.
      • Hernandez L.
      • Nelson C.
      • Lean I.
      • Santos J.
      Effects of prepartum dietary cation-anion difference and source of vitamin D in dairy cows: Lactation performance and energy metabolism.
      ). Additional data supported a trend for increased colostrum yield without an effect on colostrum components when cows were fed negative DCAD diets and cholecalciferol was replaced with calcidiol at either 1 or 3 mg/d (
      • Poindexter M.B.
      • Zimpel R.
      • Vieira-Neto A.
      • Husnain A.
      • Silva A.C.M.
      • Faccenda A.
      • Sanches de Avila A.
      • Celi P.
      • Cortinhas C.
      • Santos J.E.P.
      • Nelson C.D.
      Effect of prepartum source and amount of vitamin D supplementation on lactation performance of dairy cows.
      ) and when 0.625 mg/d of cholecalciferol was replaced with 3 mg/d of calcidiol (
      • Silva A.S.
      • Cortinhas C.S.
      • Acedo T.S.
      • Morenz M.J.F.
      • Lopes F.C.F.
      • Arrigoni M.B.
      • Ferreira M.H.
      • Jaguaribe T.L.
      • Ferreira L.D.
      • Gouvêa V.N.
      • Pereira L.G.R.
      Effects of feeding 25-hydroxyvitamin D3 with an acidogenic diet during the prepartum period in dairy cows: Mineral metabolism, energy balance, and lactation performance of Holstein dairy cows.
      ). While it remains unclear how calcidiol affected colostrum yield,
      • Martinez N.
      • Rodney R.
      • Block E.
      • Hernandez L.
      • Nelson C.
      • Lean I.
      • Santos J.
      Effects of prepartum dietary cation-anion difference and source of vitamin D in dairy cows: Lactation performance and energy metabolism.
      hypothesized calcidiol might have direct effects on epithelial cell proliferation through hormonal control or calcium and substrate availability. Colostrum IgG concentration was not affected by source of vitamin D in a study by
      • Poindexter M.B.
      • Zimpel R.
      • Vieira-Neto A.
      • Husnain A.
      • Silva A.C.M.
      • Faccenda A.
      • Sanches de Avila A.
      • Celi P.
      • Cortinhas C.
      • Santos J.E.P.
      • Nelson C.D.
      Effect of prepartum source and amount of vitamin D supplementation on lactation performance of dairy cows.
      in contrast to
      • Martinez N.
      • Rodney R.
      • Block E.
      • Hernandez L.
      • Nelson C.
      • Lean I.
      • Santos J.
      Effects of prepartum dietary cation-anion difference and source of vitamin D in dairy cows: Lactation performance and energy metabolism.
      . Further, source of vitamin D did not affect yield or composition of transition milk, collected as the second milking after calving (
      • Poindexter M.B.
      • Zimpel R.
      • Vieira-Neto A.
      • Husnain A.
      • Silva A.C.M.
      • Faccenda A.
      • Sanches de Avila A.
      • Celi P.
      • Cortinhas C.
      • Santos J.E.P.
      • Nelson C.D.
      Effect of prepartum source and amount of vitamin D supplementation on lactation performance of dairy cows.
      ). Because of the interaction between DCAD and source of vitamin D on colostral components and the inconsistent results on IgG concentration when replacing cholecalciferol with calcidiol, further research is needed to determine the mechanism of action as well as increase the external validity of previous findings.
      In addition to vitamin D, researchers have explored the source and inclusion of dietary minerals and vitamins on colostrum production. Because of limited placental transfer, certain colostral mineral and vitamin concentrations have also been considered as an important source for the newborn calf (
      • Quigley Iii J.
      • Drewry J.
      Nutrient and immunity transfer from cow to calf pre-and postcalving.
      ,
      • Przybylska J.
      • Albera E.
      • Kankofer M.
      Antioxidants in bovine colostrum.
      ). Apart from a greater IgG concentration reported by
      • Kincaid R.
      • Socha M.
      Inorganic versus complexed trace mineral supplements on performance of dairy cows.
      and
      • Formigoni A.
      • Fustini M.
      • Archetti L.
      • Emanuele S.
      • Sniffen C.
      • Biagi G.
      Effects of an organic source of copper, manganese and zinc on dairy cattle productive performance, health status and fertility.
      , replacing inorganic with organic trace minerals did not affect colostrum yield, IgG, or component concentrations (
      • Karkoodi K.
      • Chamani M.
      • Beheshti M.
      • Mirghaffari S.S.
      • Azarfar A.
      Effect of organic zinc, manganese, copper, and selenium chelates on colostrum production and reproductive and lameness indices in adequately supplemented Holstein cows.
      ,
      • Roshanzamir H.
      • Rezaei J.
      • Fazaeli H.
      Colostrum and milk performance, and blood immunity indices and minerals of Holstein cows receiving organic Mn, Zn and Cu sources.
      ,
      • Kerwin A.
      • Graef G.
      • Ryan C.
      • Ferro L.
      • Puga S.O.
      • Westhoff T.
      • Barbano D.
      • Kleinschmit D.
      • Overton T.
      Effect of replacing a portion of inorganic chloride trace minerals with trace mineral amino acid complexes.
      ,
      • Ogilvie L.
      • Van Winters B.
      • Mion B.
      • King K.
      • Spricigo J.F.W.
      • Karrow N.A.
      • Steele M.A.
      • Ribeiro E.S.
      Effects of replacing inorganic salts of trace minerals with organic trace minerals in the diet of prepartum cows on quality of colostrum and immunity of newborn calves.
      ). Further, colostrum mineral concentrations did not differ when replacing 50% of inorganic Cu, Mn, and Zn with organic proteinate sources (
      • Formigoni A.
      • Fustini M.
      • Archetti L.
      • Emanuele S.
      • Sniffen C.
      • Biagi G.
      Effects of an organic source of copper, manganese and zinc on dairy cattle productive performance, health status and fertility.
      ); however, inclusion of selenized yeast or selenium biofortified alfalfa increased Se concentrations in colostrum from Jersey and Holstein cows (
      • Weiss W.P.
      • Hogan J.S.
      Effect of selenium source on selenium status, neutrophil function, and response to intramammary endotoxin challenge of dairy cows.
      ,
      • Jaaf S.
      • Batty B.
      • Krueger A.
      • Estill C.T.
      • Bionaz M.
      Selenium biofortified alfalfa hay fed in low quantities improves selenium status and glutathione peroxidase activity in transition dairy cows and their calves.
      ,
      • Ogilvie L.
      • Van Winters B.
      • Mion B.
      • King K.
      • Spricigo J.F.W.
      • Karrow N.A.
      • Steele M.A.
      • Ribeiro E.S.
      Effects of replacing inorganic salts of trace minerals with organic trace minerals in the diet of prepartum cows on quality of colostrum and immunity of newborn calves.
      ). Although dietary supplementation of 0.2 ppm Se with 70 IU/kg vitamin E in combination with an injection of 50 mg Se and 300 IU vitamin E at 21 d before expected calving increased colostrum concentrations of α-tocopherol (
      • Weiss W.
      • Todhunter D.
      • Hogan J.
      • Smith K.
      Effect of duration of supplementation of selenium and vitamin E on periparturient dairy cows.
      ), dietary supplementation alone does not appear to be an effective strategy to increase colostral α-tocopherol concentrations (
      • Weiss W.P.
      • Hogan J.S.
      • Smith K.L.
      • Todhunter D.A.
      • Williams S.N.
      Effect of supplementing periparturient cows with vitamin E on distribution of alpha-tocopherol in blood.
      ). Similarly, maternal supplementation of 700 to 800 mg/d of β-carotene did not affect colostrum yield or concentrations of IgG, retinol, α-tocopherol, or components and yielded mixed results on colostral β-carotene concentrations (
      • Aragona K.
      • Rice E.
      • Engstrom M.
      • Erickson P.
      Effect of β-carotene supplementation to prepartum Holstein cows on colostrum quality and calf performance.
      ,
      • Prom C.M.
      • Engstrom M.A.
      • Drackley J.K.
      Effects of prepartum supplementation of &#x3b2;-carotene on colostrum and calves.
      ). Supplemental B vitamins, including nicotinic acid, increased colostrum IgG concentration (
      • Aragona K.M.
      • Chapman C.E.
      • Pereira A.B.
      • Isenberg B.
      • Standish R.
      • Maugeri C.
      • Cabral R.G.
      • Erickson P.S.
      Prepartum supplementation of nicotinic acid: Effects on health of the dam, colostrum quality, and acquisition of immunity in the calf.
      ,
      • Aragona K.
      • Rice E.
      • Engstrom M.
      • Erickson P.
      Supplementation of nicotinic acid to prepartum Holstein cows increases colostral immunoglobulin G, excretion of urinary purine derivatives, and feed efficiency in calves.
      ) while biotin altered fatty acid composition in colostrum (
      • Duplessis M.
      • Gervais R.
      • Lapierre H.
      • Girard C.L.
      Combined biotin, folic acid, and vitamin B12 supplementation given during the transition period to dairy cows: Part II. Effects on energy balance and fatty acid composition of colostrum and milk.
      ).

      Feed additives

      A limited number of feed additives have been investigated for their effect on colostrum production and results have been mixed or inconsistent. Inclusion of magnesium butyrate supplemented at 105 g/d increased colostrum yield and total IgG mass but IgG concentration and colostral components were not affected (
      • Kovács L.
      • Pajor F.
      • Bakony M.
      • Fébel H.
      • Edwards J.E.
      Prepartum Magnesium Butyrate Supplementation of Dairy Cows Improves Colostrum Yield, Calving Ease, Fertility, Early Lactation Performance and Neonatal Vitality.
      ). Feeding monensin at 24.2 g/t of total dietary DM did not affect colostrum yield or IgG concentrations (
      • Vasquez J.
      • McCarthy M.
      • Richards B.
      • Perfield K.
      • Carlson D.
      • Lock A.
      • Drackley J.
      Effects of prepartum diets varying in dietary energy density and monensin on early-lactation performance in dairy cows.
      ). Further, supplementing prepartum cows with 13.6 and 20.4 g/d of choline ions did not affect Brix % or component concentrations but
      • Swartz T.H.
      • Bradford B.J.
      • Malysheva O.
      • Caudill M.A.
      • Mamedova L.K.
      • Estes K.A.
      Effects of dietary rumen-protected choline supplementation on colostrum yields, quality, and choline metabolites from dairy cattle.
      reported a 2.9 ± 0.8 and 2.5 ± 0.8 kg increase in yield, respectively. In another study where cows were supplemented either with 0, 15, or 22 g/d of choline ions, colostrum yield was only different in the 15 g/d group, showing an increase from 3.4 (2.2–5.1) kg in control to 4.4 (2.8–6.7) kg or 5.4 (3.5–7.9) kg in cows fed 15 g/d of 2 choline products, respectively (
      • Holdorf H.T.
      • Kendall S.J.
      • Ruh K.E.
      • Caputo M.J.
      • Combs G.J.
      • Henisz S.J.
      • Brown W.E.
      • Bresolin T.
      • Ferreira R.E.P.
      • Dorea J.R.R.
      • White H.M.
      Increasing the prepartum dose of rumen-protected choline: Effects on milk production and metabolism in high-producing Holstein dairy cows.
      ). In contrast to the aforementioned studies,
      • Zenobi M.G.
      • Gardinal R.
      • Zuniga J.E.
      • Dias A.L.G.
      • Nelson C.D.
      • Driver J.P.
      • Barton B.A.
      • Santos J.E.P.
      • Staples C.R.
      Effects of supplementation with ruminally protected choline on performance of multiparous Holstein cows did not depend upon prepartum caloric intake.
      and
      • Bollatti J.
      • Zenobi M.
      • Artusso N.
      • Alfaro G.
      • Lopez A.
      • Barton B.
      • Nelson C.
      • Staples C.
      • Santos J.
      Timing of initiation and duration of feeding rumen-protected choline affects performance of lactating Holstein cows.
      did not observe an effect on colostrum yield when supplementing 12.9 g/d of choline ions. Moreover, in a field study consisting of 21 prepartum pens (n = 2,171 cows), colostrum yield as well as IgG and component concentrations were not affected by choline supplementation at 12.9 g/d (
      • Poindexter M.
      Use of Vitamin D and choline to improve health and productivy of dairy cows and their offspring. in Department of Animal Science.
      ). Including mannan oligosaccharides (
      • Franklin S.
      • Newman M.
      • Newman K.
      • Meek K.
      Immune parameters of dry cows fed mannan oligosaccharide and subsequent transfer of immunity to calves.
      ,
      • Westland A.
      • Martin R.
      • White R.
      • Martin J.
      Mannan oligosaccharide prepartum supplementation: effects on dairy cow colostrum quality and quantity.
      ) or direct-fed microbials and enzymes (
      • Ort S.B.
      • Aragona K.M.
      • Chapman C.E.
      • Shangraw E.
      • Brito A.
      • Schauff D.
      • Erickson P.S.
      The impact of direct-fed microbials and enzymes on the health and performance of dairy cows with emphasis on colostrum quality and serum immunoglobulin concentrations in calves.
      ,
      • Real L.
      Effects of Native Rumen Microbes as a DFM on Colostrum in Holstein Dairy Cows.
      ) in the prepartum diet has yielded mixed results on colostrum production. Contrary to
      • Franklin S.
      • Newman M.
      • Newman K.
      • Meek K.
      Immune parameters of dry cows fed mannan oligosaccharide and subsequent transfer of immunity to calves.
      and
      • Ort S.B.
      • Aragona K.M.
      • Chapman C.E.
      • Shangraw E.
      • Brito A.
      • Schauff D.
      • Erickson P.S.
      The impact of direct-fed microbials and enzymes on the health and performance of dairy cows with emphasis on colostrum quality and serum immunoglobulin concentrations in calves.
      , colostrum yield was increased by supplementing 2 g/d of Saccharomyces cerevisiae derived mannan oligosaccharides (
      • Westland A.
      • Martin R.
      • White R.
      • Martin J.
      Mannan oligosaccharide prepartum supplementation: effects on dairy cow colostrum quality and quantity.
      ) or 5 g/d of direct-fed microbials and enzymes (
      • Real L.
      Effects of Native Rumen Microbes as a DFM on Colostrum in Holstein Dairy Cows.
      ). The influence of feed additives and supplementation requires further consideration and replication of studies before conclusions can be drawn for on-farm use.

      Prepartum Management

      Prepartum environment

      The seasonality of colostrum production, as described above, is confounded by changes in environmental conditions including exposure to light as well as heat exposure, measured as temperature-humidity index (THI). Authors have associated THI or photoperiod with colostrum yield (
      • Cabral R.G.
      • Chapman C.E.
      • Aragona K.M.
      • Clark E.
      • Lunak M.
      • Erickson P.S.
      Predicting colostrum quality from performance in the previous lactation and environmental changes.
      ,
      • Gavin K.
      • Neibergs H.
      • Hoffman A.
      • Kiser J.N.
      • Cornmesser M.A.
      • Haredasht S.A.
      • Martinez-Lopez B.
      • Wenz J.R.
      • Moore D.A.
      Low colostrum yield in Jersey cattle and potential risk factors.
      ,
      • Borelli E.
      • Jonsson N.N.
      • Denholm K.S.
      Associations between putative risk factors and poor colostrum yield in Holstein Friesian cattle.
      ,
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      ), IgG concentrations or Brix % (
      • Cabral R.G.
      • Chapman C.E.
      • Aragona K.M.
      • Clark E.
      • Lunak M.
      • Erickson P.S.
      Predicting colostrum quality from performance in the previous lactation and environmental changes.
      ,
      • Shivley C.B.
      • Lombard J.E.
      • Urie N.J.
      • Haines D.M.
      • Sargent R.
      • Kopral C.A.
      • Earleywine T.J.
      • Olson J.D.
      • Garry F.B.
      Preweaned heifer management on US dairy operations: Part II. Factors associated with colostrum quality and passive transfer status of dairy heifer calves.
      ,
      • Zentrich E.
      • Iwersen M.
      • Wiedrich M.C.
      • Drillich M.
      • Klein-Jöbstl D.
      Short communication: Effect of barn climate and management-related factors on bovine colostrum quality.
      ), and colostral composition (
      • Nardone A.
      • Lacetera N.
      • Bernabucci U.
      • Ronchi B.
      Composition of colostrum from dairy heifers exposed to high air temperatures during late pregnancy and the early postpartum period.
      ,
      • Román L.
      • Piñeyrúa J.T.M.
      • Banchero G.
      • La Manna A.
      Access to shade during the dry period improves the performance of multiparous Holstein cows.
      ,
      • Alward K.J.
      Photoperiod and Temperature-Humidity Index during the Dry-Period Impact Colostrum and Milk Production in Dairy Cattle. in Department of Animal Sciences.
      ). When THI 7-d before calving and light intensity 14-d before calving were categorized, colostrum yield from multiparous Holstein cows increased as THI and light intensity categories increased (
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      ).
      • Gavin K.
      • Neibergs H.
      • Hoffman A.
      • Kiser J.N.
      • Cornmesser M.A.
      • Haredasht S.A.
      • Martinez-Lopez B.
      • Wenz J.R.
      • Moore D.A.
      Low colostrum yield in Jersey cattle and potential risk factors.
      reported a positive correlation between colostrum yield from Jersey cows and photoperiod as well as maximum weekly THI. Nevertheless, the lack of studies considering photoperiod and THI independent of one another in controlled settings makes it challenging to determine if a causal relationship exists. In a study by
      • Alward K.J.
      Photoperiod and Temperature-Humidity Index during the Dry-Period Impact Colostrum and Milk Production in Dairy Cattle. in Department of Animal Sciences.
      , colostrum yield, Brix %, and IgG concentration from Holstein and Jersey cows did not differ when cows were managed for a short or long-day photoperiod (8 vs. 16 h of light/d). Similarly, increasing photoperiod from 8 to 16 h of light/d for the entire dry period did not affect colostrum yield and IgG concentration (
      • Morin D.E.
      • Nelson S.V.
      • Reid E.D.
      • Nagy D.W.
      • Dahl G.E.
      • Constable P.D.
      Effect of colostral volume, interval between calving and first milking, and photoperiod on colostral IgG concentrations in dairy cows.
      ). Although more data are needed to determine if increased light exposure during the dry period can affect colostrum production, short day light exposure (8 h/d) during the dry period remains the optimal lighting program during the dry period for the resulting benefits in lactation performance (
      • Auchtung T.L.
      • Rius A.G.
      • Kendall P.E.
      • McFadden T.B.
      • Dahl G.E.
      Effects of photoperiod during the dry period on prolactin, prolactin receptor, and milk production of dairy cows.
      ,
      • Velasco J.M.
      • Reid E.D.
      • Fried K.K.
      • Gressley T.F.
      • Wallace R.L.
      • Dahl G.E.
      Short-day photoperiod increases milk yield in cows with a reduced dry period length.
      ).
      Regarding the effect of heat stress, heat abatement strategies appear to influence colostrum production (
      • Nardone A.
      • Lacetera N.
      • Bernabucci U.
      • Ronchi B.
      Composition of colostrum from dairy heifers exposed to high air temperatures during late pregnancy and the early postpartum period.
      ,
      • Karimi M.T.
      • Ghorbani G.R.
      • Kargar S.
      • Drackley J.K.
      Late-gestation heat stress abatement on performance and behavior of Holstein dairy cows.
      ). In a study by
      • Almoosavi S.S.
      • Ghoorchi T.
      • Naserian A.
      • Khanaki H.
      • Drackley J.
      • Ghaffari M.
      Effects of late-gestation heat stress independent of reduced feed intake on colostrum, metabolism at calving, and milk yield in early lactation of dairy cows.
      , cooled cows (access to shade, sprinklers, and fans) produced more colostrum (7.1 ± 0.6 kg) with an elevated IgG concentration (92.2 ± 2.5 g/L) compared with heat-stressed cows (access to shade but not to sprinklers or fans; 4.0 ± 0.6 kg; 74.7 ± 2.5 g/L). In addition, compared with the cooled cows, colostrum yield (6.0 ± 0.6 kg) and IgG concentration (88.5 ± 2.5 g/L) did not differ in a third group of cows that were cooled but offered the same amount of feed as the heat-stressed group (
      • Almoosavi S.S.
      • Ghoorchi T.
      • Naserian A.
      • Khanaki H.
      • Drackley J.
      • Ghaffari M.
      Effects of late-gestation heat stress independent of reduced feed intake on colostrum, metabolism at calving, and milk yield in early lactation of dairy cows.
      ). As has been shown with milk production (
      • Baumgard L.H.
      • Rhoads Jr., R.P.
      Effects of heat stress on postabsorptive metabolism and energetics.
      ), reduced feed intake only partially explains a lower colostrum yield as a result of heat stress in the aforementioned study and provides further support for thermal management of dry cows beyond the positive effects on cow and calf productivity (
      • Tao S.
      • Dahl G.
      Invited review: Heat stress effects during late gestation on dry cows and their calves.
      ,
      • Laporta J.
      • Fabris T.
      • Skibiel A.
      • Powell J.
      • Hayen M.
      • Horvath K.
      • Miller-Cushon E.
      • Dahl G.
      In utero exposure to heat stress during late gestation has prolonged effects on the activity patterns and growth of dairy calves.
      ,
      • Laporta J.
      • Ferreira F.C.
      • Ouellet V.
      • Dado-Senn B.
      • Almeida A.K.
      • De Vries A.
      • Dahl G.E.
      Late-gestation heat stress impairs daughter and granddaughter lifetime performance.
      ).

      Dry period length

      Observational data suggest that decreasing the length of the dry period results in a lower colostrum yield (
      • Poindexter M.
      Use of Vitamin D and choline to improve health and productivy of dairy cows and their offspring. in Department of Animal Science.
      ,
      • Soufleri A.
      • Banos G.
      • Panousis N.
      • Fletouris D.
      • Arsenos G.
      • Kougioumtzis A.
      • Valergakis G.E.
      Evaluation of Factors Affecting Colostrum Quality and Quantity in Holstein Dairy Cattle.
      ,
      • Borelli E.
      • Jonsson N.N.
      • Denholm K.S.
      Associations between putative risk factors and poor colostrum yield in Holstein Friesian cattle.
      ). In a study on 12,553 Holstein cows from 18 dairy farms,
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      reported an increase in colostrum yield as the length of the dry period, categorized as < 47, 47–67, or > 67-d, increased. Jersey cows with a 45-d dry period had a 1.7 times greater odds of producing < 2.7 kg of colostrum compared with cows with a 65-d dry period. Further, researchers revealed a 60-d dry period resulted in +2.2 ± 0.4 and +2.6 ± 0.6 kg more colostrum compared with cows managed for a shortened (30–40 d) dry period (
      • Grusenmeyer D.
      • Ryan C.
      • Galton D.
      • Overton T.
      Shortening the dry period from 60 to 40 days does not affect colostrum quality but decreases colostrum yield by Holstein cows.
      ,
      • Mayasari N.
      • de Vries Reilingh G.
      • Nieuwland M.G.
      • Remmelink G.J.
      • Parmentier H.K.
      • Kemp B.
      • van Knegsel A.T.
      Effect of maternal dry period length on colostrum immunoglobulin content and on natural and specific antibody titers in calves.
      ). Colostrum IgG concentration was not affected by a dry period shortened to 35 to 40 d (
      • Grusenmeyer D.
      • Ryan C.
      • Galton D.
      • Overton T.
      Shortening the dry period from 60 to 40 days does not affect colostrum quality but decreases colostrum yield by Holstein cows.
      ,
      • Cermakova J.
      • Kudrna V.
      • Simeckova M.
      • Vyborna A.
      • Dolezal P.
      • Illek J.
      Comparison of shortened and conventional dry period management strategies.
      ,
      • Shoshani E.
      • Rozen S.
      • Doekes J.J.
      Effect of a short dry period on milk yield and content, colostrum quality, fertility, and metabolic status of Holstein cows.
      ), but was reduced approximately 30–59% in the absence of a dry period compared with a shortened dry period (
      • Rastani R.
      • Grummer R.
      • Bertics S.
      • Gümen A.
      • Wiltbank M.
      • Mashek D.
      • Schwab M.
      Reducing dry period length to simplify feeding transition cows: milk production, energy balance, and metabolic profiles.
      ,
      • Klusmeyer T.
      • Fitzgerald A.
      • Fabellar A.
      • Ballam J.
      • Cady R.
      • Vicini J.
      Effect of recombinant bovine somatotropin and a shortened or no dry period on the performance of lactating dairy cows.
      ,
      • Mayasari N.
      • de Vries Reilingh G.
      • Nieuwland M.G.
      • Remmelink G.J.
      • Parmentier H.K.
      • Kemp B.
      • van Knegsel A.T.
      Effect of maternal dry period length on colostrum immunoglobulin content and on natural and specific antibody titers in calves.
      ). Additionally, a dry period ≥ 85-d was associated with greater concentrations of colostrum fat, but protein concentration was not affected by dry period length (
      • Soufleri A.
      • Banos G.
      • Panousis N.
      • Fletouris D.
      • Arsenos G.
      • Kougioumtzis A.
      • Valergakis G.E.
      Evaluation of Factors Affecting Colostrum Quality and Quantity in Holstein Dairy Cattle.
      ).

      Time in the close-up pen

      Since colostrum synthesis exerts a metabolic demand during late gestation, prepartum diet formulation (discussed above) as well as length of exposure to the close-up ration has been evaluated in 2-phase dry cow systems. Contrary to
      • Weich W.
      • Block E.
      • Litherland N.
      Extended negative dietary cation-anion difference feeding does not negatively affect postpartum performance of multiparous dairy cows.
      ,
      • Lopera C.
      • Zimpel R.
      • Vieira-Neto A.
      • Lopes F.R.
      • Ortiz W.
      • Poindexter M.
      • Faria B.N.
      • Gambarini M.L.
      • Block E.
      • Nelson C.D.
      • Santos J.E.P.
      Effects of level of dietary cation-anion difference and duration of prepartum feeding on performance and metabolism of dairy cows.
      , and
      • Sørensen M.M.
      Effects of close-up period duration and feeding level on periparturient performance and health of dairy cows. in Department of Veterinary and Animal Sciences. Vol.
      ,
      • Farahani T.A.
      • Amanlou H.
      • Kazemi-Bonchenari M.
      Effects of shortening the close-up period length coupled with increased supply of metabolizable protein on performance and metabolic status of multiparous Holstein cows.
      observed a trend for a 2.0 ± 0.8 kg greater colostrum yield with a 21-d compared with a 10-d close-up period. The aforementioned increase in yield was larger than that observed in an observational study (10 d = 4.9 ± 0.2 vs. 20 d = 5.2 ± 0.1 kg;
      • Borchardt S.
      • Sutter F.
      • Heuwieser W.
      • Venjakob P.
      Management-related factors in dry cows and their associations with colostrum quantity and quality on a large commercial dairy farm.
      ). Yet, when categorized as ≤ 15, 16–30, or > 30 d, time in the close-up pen was not associated with colostrum yield or Brix % in an analysis of 16,032 Holstein cows (
      • Westhoff T.A.
      • Womack S.J.
      • Overton T.R.
      • Ryan C.M.
      • Mann S.
      Epidemiology of bovine colostrum production in New York Holstein herds: cow, management, and environmental factors.
      ). Pen stocking density ranging from 41 to 163% was not associated with colostrum yield in an observational analysis of 238 cows (
      • Borelli E.
      • Jonsson N.N.
      • Denholm K.S.
      Associations between putative risk factors and poor colostrum yield in Holstein Friesian cattle.
      ). Further, decreasing stocking density (100% headlock, 109% stalls vs. 80% headlock, 86.3% stalls; Silva et al., 2016 and 120% vs. 100% vs. 80% headlock and stalls; Jiang et al., 2021) of the close-up pen did not affect colostrum yield or Brix percentage. While it appears that total dry period length is more influential for colostrum synthesis compared with time and stocking density of the close-up pen, recent evidence suggests that the interaction of pen move with dry cow booster vaccinations to increase specific calf-health related antibodies in colostrum should also be considered. Cows administered booster vaccinations at 28 d relative to calving and moved to the close-up pen at 21 d relative to calving had greater colostral IgG concentration (160.4 ± 7.0 g/L) compared with cows vaccinated and moved at 21 d relative to calving (134.4 ± 7.0 g/L) but neither treatment differed from cows vaccinated and moved to the close-up pen at 28 d relative to calving (148.3 ± 7.2 g/L) (
      • Menichetti B.
      • Garcia-Guerra A.
      • Lakritz J.
      • Weiss W.
      • Velez J.
      • Bothe H.
      • Merchan D.
      • Schuenemann G.
      Effects of prepartum vaccination timing relative to pen change with an acidogenic diet on serum and colostrum immunoglobulins in Holstein dairy cows.
      ). Although replication of these data is lacking, results of this study suggest that vaccination one week earlier and not coinciding with potential detrimental effects of a pen move was more beneficial than earlier vaccination in combination with a pen move. When timing of vaccination is considered independent of pen moves, administration of vaccines earlier in the dry period or given repeatedly, and consistent with manufacturer label recommendations, might offer a management strategy to increase vaccine derived antibodies in colostrum since most colostral immunoglobulins are produced in and transferred from maternal circulation.

      Dry off procedure and udder health

      Selective and blanket dry cow therapy protocols are commonly used to reduce intramammary infection (
      • Zwald A.
      • Ruegg P.
      • Kaneene J.
      • Warnick L.
      • Wells S.
      • Fossler C.
      • Halbert L.
      Management practices and reported antimicrobial usage on conventional and organic dairy farms.
      ,
      • Winder C.
      • Sargeant J.
      • Kelton D.
      • Leblanc S.
      • Duffield T.
      • Glanville J.
      • Wood H.
      • Churchill K.
      • Dunn J.
      • Bergevin M.
      • Dawkins K.
      • Meadows S.
      • O’Connor A.M.
      Comparative efficacy of blanket versus selective dry-cow therapy: A systematic review and pairwise meta-analysis.
      ), but limited data are available on their effect on colostrum production. In studies using a teat-sealant only or an antibiotic in addition to a teat sealant on cows with a low risk (SCC < 200,000 cells/mL) for intramammary infection,
      • Lavery A.
      • Craig A.-L.
      • Gordon A.W.
      • Ferris C.P.
      Impact of adopting non-antibiotic dry-cow therapy on the performance and udder health of dairy cows.
      and
      • Vasquez A.
      • Nydam D.
      • Foditsch C.
      • Warnick L.
      • Wolfe C.
      • Doster E.
      • Morley P.S.
      Characterization and comparison of the microbiomes and resistomes of colostrum from selectively treated dry cows.
      found no differences in colostrum fat, protein, lactose, and IgG concentrations as well as the colostrum microbiome. However, the effect of dry-cow therapy might depend on the risk of intramammary infections as the yield of colostrum from a persistently infected gland, defined as growth of ≥ 50 cfu/mL of the respective mastitis-causing pathogen when sampled 14 and 7 d relative to calving, was reduced compared with non-infected glands (
      • Maunsell F.
      • Morin D.
      • Constable P.D.
      • Hurley W.
      • McCoy G.
      • Kakoma I.
      • Isaacson R.
      Effects of mastitis on the volume and composition of colostrum produced by Holstein cows.
      ). Although clinical mastitis in the previous lactation was not associated with colostrum yield, Brix %, or IgG concentrations (
      • Aghakhani M.
      • Shahraki A.D.F.
      • Tabatabaei S.
      • Toghyani M.
      • Rafiee H.
      Cow-level factors associated with colostrum yield and quality of Holstein dairy cows.
      ) and IgG, IgM, and IgA as well as fat, protein, and lactose concentration were not affected by intramammary infection (
      • Maunsell F.
      • Morin D.
      • Constable P.D.
      • Hurley W.
      • McCoy G.
      • Kakoma I.
      • Isaacson R.
      Effects of mastitis on the volume and composition of colostrum produced by Holstein cows.
      ,
      • Enger K.
      • Hardy N.
      • Enger B.
      Relationship between intramammary infection and antibody concentrations in Jersey and Holstein colostrum.
      ,
      • Pikhtirova A.
      • Pecka-Kiełb E.
      • Zachwieja A.
      • Bujok J.
      • Zigo F.
      Effects of non-aureus staphylococci on colostrum composition, properties and fatty acid profile in cow–a preliminary study.
      ), maintaining udder health during the dry period should remain a priority independent of considerations for colostrum production.

      HARVESTING COLOSTRUM

      Time to Colostrum Harvest

      The time near parturition marks the transition from lactogenesis I, including colostrum synthesis, to copious milk production (lactogenesis II;
      • Baumrucker C.R.
      • Bruckmaier R.M.
      Colostrogenesis: IgG1 transcytosis mechanisms.
      ) . Concentrations of most colostral components and bioactive factors (fat, protein, total solids, SCC, immunoglobulins, oligosaccharides, miRNA, insulin, IGF-1, minerals, vitamins, etc.) decrease in subsequent hours to days following parturition while milk yield and lactose concentrations increase (
      • Foley J.
      • Otterby D.
      Availability, storage, treatment, composition, and feeding value of surplus colostrum: a review.
      ,
      • Blum J.
      • Hammon H.
      Colostrum effects on the gastrointestinal tract, and on nutritional, endocrine and metabolic parameters in neonatal calves.
      ,
      • Hammon H.M.
      • Zanker I.A.
      • Blum J.W.
      Delayed colostrum feeding affects IGF-I and insulin plasma concentrations in neonatal calves.
      ,
      • Fahey M.
      • Fischer A.
      • Steele M.
      • Greenwood S.
      Characterization of the colostrum and transition milk proteomes from primiparous and multiparous Holstein dairy cows.
      ,
      • Wilms J.
      • Hare K.
      • Fischer-Tlustos A.
      • Vahmani P.
      • Dugan M.
      • Leal L.
      • Steele M.
      Fatty acid profile characterization in colostrum, transition milk, and mature milk of primi-and multiparous cows during the first week of lactation.
      ). As such, timely colostrum harvest has been recommended.
      • Quigley J.D.
      • Lago A.
      • Chapman C.
      • Erickson P.
      • Polo J.
      Evaluation of the Brix refractometer to estimate immunoglobulin G concentration in bovine colostrum.
      observed a quadratic relationship between IgG concentration and time from calving to colostrum harvest such that IgG concentration was lower when collected ≥ 8 h post calving. In agreement, other authors have observed decreased colostrum Brix % or IgG concentration in colostrum harvested ≥6 to 9 h post-calving and an increased colostrum yield when harvested ≥12 h post-calving (
      • Conneely M.
      • Berry D.P.
      • Sayers R.
      • Murphy J.P.
      • Lorenz I.
      • Doherty M.L.
      • Kennedy E.
      Factors associated with the concentration of immunoglobulin G in the colostrum of dairy cows.
      ,
      • Silva-del-Río N.
      • Rolle D.
      • García-Muñoz A.
      • Rodríguez-Jiménez S.
      • Valldecabres A.
      • Lago A.
      • Pandey P.
      Colostrum immunoglobulin G concentration of multiparous Jersey cows at first and second milking is associated with parity, colostrum yield, and time of first milking, and can be estimated with Brix refractometry.
      ,
      • Soufleri A.
      • Banos G.
      • Panousis N.
      • Fletouris D.
      • Arsenos G.
      • Kougioumtzis A.
      • Valergakis G.E.
      Evaluation of Factors Affecting Colostrum Quality and Quantity in Holstein Dairy Cattle.
      ). To maximize colostral components and IgG concentration, we recommend harvesting colostrum ≤ 8 h following calving. Notably, producers that feed a calf its dam's colostrum should prioritize colostrum harvest ≤ 2 h from calving to ensure timely ingestion of colostrum for the newborn calf.

      Administration of Oxytocin

      As with milk letdown, release of oxytocin from the pituitary gland is a necessary response for a complete colostrum harvest and is most often achieved by tactile teat stimulation (
      • Bruckmaier R.
      • Blum J.
      Oxytocin release and milk removal in ruminants.
      ). Therefore, disruptions in oxytocin release, as has been reported in primiparous cows and cows milked in unfamiliar locations (
      • Bruckmaier R.M.
      • Schams D.
      • Blum J.W.
      Aetiology of disturbed milk ejection in parturient primiparous cows.
      ,
      • Bruckmaier R.M.
      • Schams D.
      • Blum J.W.
      Milk removal in familiar and unfamiliar surroundings: concentrations of oxytocin, prolactin, cortisol and β–endorphin.
      ) might prevent a complete removal of colostrum. Contrary to the study hypothesis,
      • Sutter F.
      • Borchardt S.
      • Schuenemann G.M.
      • Rauch E.
      • Erhard M.
      • Heuwieser W.
      Evaluation of 2 different treatment procedures after calving to improve harvesting of high-quantity and high-quality colostrum.
      found that colostrum yield was not affected when the calf was present before and during colostrum harvest or when administering 20 IU of oxytocin intramuscularly 3 min. before manual stimulation in preparation for colostrum harvest, but IgG concentration was increased by 5.3 ± 2.6 and 6.3 ± 2.7 g/L, respectively. Elevated oxytocin concentrations can alter the tight junctions of the mammary gland (
      • Allen J.
      Milk synthesis and secretion rates in cows with milk composition changed by oxytocin.
      ,
      • Wall S.K.
      • Wellnitz O.
      • Hernández-Castellano L.E.
      • Ahmadpour A.
      • Bruckmaier R.M.
      Supraphysiological oxytocin increases the transfer of immunoglobulins and other blood components to milk during lipopolysaccharide-and lipoteichoic acid–induced mastitis in dairy cows.
      ,
      • Farmer C.
      • Lessard M.
      • Knight C.
      • Quesnel H.
      Oxytocin injections in the postpartal period affect mammary tight junctions in sows.
      ) which might have affected IgG concentration in the above-mentioned study, leading to the small observed effect. Oxytocin injections have not been associated with changes in fat, protein, or lactose concentrations in milk (
      • Nostrand S.
      • Galton D.
      • Erb H.
      • Bauman D.
      Effects of daily exogenous oxytocin on lactation milk yield and composition.
      ,
      • Ballou L.U.
      • Bleck J.L.
      • Bleck G.T.
      • Bremel R.D.
      The effects of daily oxytocin injections before and after milking on milk production, milk plasmin, and milk composition.
      ) although the effect of oxytocin administration or presence of the calf on other colostral components has not been reported to the knowledge of the authors. Given findings reported by
      • Sutter F.
      • Borchardt S.
      • Schuenemann G.M.
      • Rauch E.
      • Erhard M.
      • Heuwieser W.
      Evaluation of 2 different treatment procedures after calving to improve harvesting of high-quantity and high-quality colostrum.
      originated from a single commercial dairy farm, the external validity on these data remain uncertain, and create a need for additional studies on multiple dairy farms with varying widely used premilking routines. Further, research is needed to determine whether oxytocin administration at colostrum harvest affects the milk letdown reflex at subsequent milkings.

      POST-HARVEST COLOSTRUM MANAGEMENT

      On-farm Assessment of Colostrum Quality

      Assays to determine colostral IgG [radial immunodiffusion (RID), enzyme-linked immunosorbent assay (ELISA), and turbidimetric immune assay (TIA)] are time-consuming and costly, making them infeasible for commercial dairy producers. Moreover, because of a bias between ELISA and TIA when compared with RID (
      • Gelsinger S.L.
      • Smith A.M.
      • Jones C.M.
      • Heinrichs A.J.
      Technical note: Comparison of radial immunodiffusion and ELISA for quantification of bovine immunoglobulin G in colostrum and plasma.
      ,
      • Dunn A.
      • Duffy C.
      • Gordon A.
      • Morrison S.
      • Argűello A.
      • Welsh M.
      • Earley B.
      Comparison of single radial immunodiffusion and ELISA for the quantification of immunoglobulin G in bovine colostrum, milk and calf sera.
      ,
      • Breuer R.M.
      • Wiley C.
      • Dohlman T.
      • Smith J.S.
      • McKeen L.
      • Kreuder A.J.
      Comparison of turbidometric immunoassay and brix refractometry to radial immunodiffusion for assessment of colostral immunoglobulin concentration in beef cattle.
      ,
      • Röder M.
      • Borchardt S.
      • Heuwieser W.
      • Rauch E.
      • Sargent R.
      • Sutter F.
      Evaluation of an ELISA and four on-farm tests to estimate colostrum quality for dairy cows.
      ), comparing results between methods is not recommended. However, determining the specific gravity or refractive index of colostrum via a hydrometer and Brix refractometer, respectively, have been investigated for their role as rapid and affordable indirect estimates of colostrum quality. When compared with IgG determined by RID, a hydrometer and Brix refractometer exhibited a moderate to strong correlation (hydrometer: r = 0.58 to 0.79; Brix refractometer: r = 0.64 to 0.75;
      • Bielmann V.
      • Gillan J.
      • Perkins N.R.
      • Skidmore A.L.
      • Godden S.
      • Leslie K.E.
      An evaluation of Brix refractometry instruments for measurement of colostrum quality in dairy cattle.
      ,
      • Quigley J.D.
      • Lago A.
      • Chapman C.
      • Erickson P.
      • Polo J.
      Evaluation of the Brix refractometer to estimate immunoglobulin G concentration in bovine colostrum.
      ,
      • Bartier A.L.
      • Windeyer M.C.
      • Doepel L.
      Evaluation of on-farm tools for colostrum quality measurement.
      ,
      • Morrill K.
      • Robertson K.
      • Spring M.
      • Robinson A.
      • Tyler H.
      Validating a refractometer to evaluate immunoglobulin G concentration in Jersey colostrum and the effect of multiple freeze–thaw cycles on evaluating colostrum quality.
      ,
      • Röder M.
      • Borchardt S.
      • Heuwieser W.
      • Rauch E.
      • Sargent R.
      • Sutter F.
      Evaluation of an ELISA and four on-farm tests to estimate colostrum quality for dairy cows.
      ). Of note, colostrum temperature as well as composition can affect the specific gravity of colostrum (
      • Mechor G.
      • Gröhn Y.
      • McDowell L.
      • Van Saun R.
      Specific gravity of bovine colostrum immunoglobulins as affected by temperature and colostrum components.
      ,
      • Morin D.E.
      • Constable P.D.
      • Maunsell F.P.
      • McCoy G.C.
      Factors associated with colostral specific gravity in dairy cows.
      ). Using a cut-point of ≥ 22.0%, a meta-analysis revealed the posttest probability of an IgG concentration ≥ 50.0 g/L was 94.3% (95% CI: 90.7–96.9%) (
      • Buczinski S.
      • Vandeweerd J.
      Diagnostic accuracy of refractometry for assessing bovine colostrum quality: A systematic review and meta-analysis.
      ). For the hydrometer, the negative predictive value (probability of a hydrometer result to correctly identify a sample as ≥ 50.0 g of IgG/L) with a cut point of 1,047 was 97.1 (95% CI: 92.8–99.2%;
      • Röder M.
      • Borchardt S.
      • Heuwieser W.
      • Rauch E.
      • Sargent R.
      • Sutter F.
      Evaluation of an ELISA and four on-farm tests to estimate colostrum quality for dairy cows.
      ) suggesting both hydrometers and Brix refractometers offer suitable in-direct estimates to identify high quality colostrum for on-farm use. In a study by
      • Godden S.M.
      • Hazel A.
      Relationship between milking fraction and immunoglobulin G concentration in first milking colostrum from holstein cows.
      , colostrum collected at the beginning of the milking process resulted in a higher IgG concentration compared with a composite sample and to samples collected during the milking process. Collection of a composite sample from the bucket after the milking process is recommended for quality assessment. Although high-quality colostrum is currently defined as an IgG concentration ≥ 50 g/L, recommended IgG intake through colostrum might change as our understanding of the short-term and long-term benefits of achieving higher concentrations of IgG in the calf rather than merely surpassing a minimum threshold grows (
      • Lombard J.
      • Urie N.
      • Garry F.
      • Godden S.
      • Quigley J.
      • Earleywine T.
      • McGuirk S.
      • Moore D.
      • Branan M.
      • Chamorro M.
      • Smith G.
      • Shivley C.
      • Catherman D.
      • Haines D.
      • Heinrichs A.J.
      • James R.
      • Maas J.
      • Sterner K.
      Consensus recommendations on calf-and herd-level passive immunity in dairy calves in the United States.
      ).

      Bacterial and Pathogen Contamination

      Along with preserving the nutritional and bioactive components, minimizing contamination is fundamental to successful colostrum management. Feeding contaminated colostrum can reduce absorption of immunoglobulins (
      • James R.
      • Polan C.
      • Cummins K.
      Influence of administered indigenous microorganisms on uptake of [iodine-125] γ-globulin in vivo by intestinal segments of neonatal calves.
      ,
      • Elizondo-Salazar J.
      • Heinrichs A.J.
      Feeding heat-treated colostrum or unheated colostrum with two different bacterial concentrations to neonatal dairy calves.
      ) as well as expose calves to pathogenic microorganisms (
      • Streeter R.N.
      • Hoffsis G.
      • Bech-Nielsen S.
      • Shulaw W.
      • Rings D.
      Isolation of Mycobacterium paratuberculosis from colostrum and milk of subclinically infected cows.
      ,
      • Godden S.
      • McMartin S.
      • Feirtag J.
      • Stabel J.
      • Bey R.
      • Goyal S.
      • Metzger L.
      • Fetrow J.
      • Wells S.
      • Chester-Jones H.
      Heat-treatment of bovine colostrum. II: effects of heating duration on pathogen viability and immunoglobulin G.
      ). However, minimizing bacterial contamination and pathogen transfer via colostrum remains an area of opportunity. In an evaluation of 1,241 colostrum samples from 39 Czech farms, only 352 (28.4%) and 1,095 (88.2%) samples were below the industry standard total plate count (<100,000 cfu/mL;
      • McGuirk S.M.
      • Collins M.
      Managing the production, storage, and delivery of colostrum.
      ) and total coliform count (<10,000 cfu/mL;
      • McGuirk S.M.
      • Collins M.
      Managing the production, storage, and delivery of colostrum.
      ), respectively (
      • Šlosárková S.
      • Pechová A.
      • Staněk S.
      • Fleischer P.
      • Zouharová M.
      • Nejedlá E.
      Microbial contamination of harvested colostrum on Czech dairy farms.
      ). Further,
      • Morrill K.M.
      • Conrad E.
      • Lago A.
      • Campbell J.
      • Quigley J.
      • Tyler H.
      Nationwide evaluation of quality and composition of colostrum on dairy farms in the United States.
      noted that 409 of 746 (54.8%) colostrum samples from 67 farms in the United States were below the industry standard total plate count. As we further understand the effects that bacterial and pathogen contamination have on the calf, industry standard thresholds for total plate and coliform counts among other contaminants might arise or need to be reevaluated. Although some pathogens can shed in the mammary gland, significant pathogen and environmental contamination occur particularly during harvest but also during storage or feeding of colostrum (
      • Stewart S.
      • Godden S.
      • Bey R.
      • Rapnicki P.
      • Fetrow J.
      • Farnsworth R.
      • Scanlon M.
      • Arnold Y.
      • Clow L.
      • Mueller K.
      • Ferrouillet C.
      Preventing bacterial contamination and proliferation during the harvest, storage, and feeding of fresh bovine colostrum.
      ). In fact, of the 155 cultured colostrum samples, 21 (13.5%) samples were positive for a gram-positive mastitis agent while 117 (75.5%), 127 (81.9%), and 128 (82.6%) samples resulted in environmental, fecal, and skin and mucosa contaminants, respectively (
      • Šlosárková S.
      • Pechová A.
      • Staněk S.
      • Fleischer P.
      • Zouharová M.
      • Nejedlá E.
      Microbial contamination of harvested colostrum on Czech dairy farms.
      ). As such, special consideration should be given to identify the risk of pathogen shedding in the mammary gland as well as minimize fecal and environmental contamination (
      • Streeter R.N.
      • Hoffsis G.
      • Bech-Nielsen S.
      • Shulaw W.
      • Rings D.
      Isolation of Mycobacterium paratuberculosis from colostrum and milk of subclinically infected cows.
      ,
      • Stabel J.R.
      On-Farm Batch Pasteurization Destroys Mycobacterium paratuberculosis in Waste Milk.
      ). In addition, cleaning and sanitizing all equipment that comes in contact with the colostrum (
      • Renaud D.L.
      • Kelton D.F.
      • LeBlanc S.J.
      • Haley D.B.
      • Jalbert A.B.
      • Duffield T.F.
      Validation of commercial luminometry swabs for total bacteria and coliform counts in colostrum-feeding equipment.
      ,
      • Buczinski S.
      • Morin M.-P.
      • Roy J.-P.
      • Rousseau M.
      • Villettaz-Robichaud M.
      • Dubuc J.
      Use of ATP luminometry to assess the cleanliness of equipment used to collect and feed colostrum on dairy farms.
      ) as well as rapidly cooling colostrum (using ice or cold water) after harvest and properly treating and storing colostrum can aid in reducing bacterial replication and preserving the nutritional and bioactive components.

      Heat Treatment

      Heat-treatment can be an effective strategy to decrease total bacterial counts in colostrum with minimal effect on immunoglobulin G concentration when treated at 60°C for 60 min. (
      • Godden S.
      • McMartin S.
      • Feirtag J.
      • Stabel J.
      • Bey R.
      • Goyal S.
      • Metzger L.
      • Fetrow J.
      • Wells S.
      • Chester-Jones H.
      Heat-treatment of bovine colostrum. II: effects of heating duration on pathogen viability and immunoglobulin G.
      ,
      • Elizondo-Salazar J.A.
      • Jayarao B.M.
      • Heinrichs A.J.
      Effect of heat treatment of bovine colostrum on bacterial counts, viscosity, and Immunoglobulin G concentration.
      ,
      • Donahue M.
      • Godden S.M.
      • Bey R.
      • Wells S.
      • Oakes J.M.
      • Sreevatsan S.
      • Stabel J.
      • Fetrow J.
      Heat treatment of colostrum on commercial dairy farms decreases colostrum microbial counts while maintaining colostrum immunoglobulin G concentrations.
      ,
      • Hesami S.
      • Shahraki A.-D.F.
      • Zakian A.
      • Ghalamkari G.
      The effect of various heat-treatment methods on colostrum quality, health and performance of dairy calves.
      ,
      • Malik M.I.
      • Rashid M.A.
      • Raboisson D.
      Heat treatment of colostrum at 60 degrees C decreases colostrum immunoglobulins but increases serum immunoglobulins and serum total protein: A meta-analysis.
      ). In a recent meta-analysis, the loss of IgG when colostrum was heat treated at ≤ 60°C and 60–63°C was −3.6 (−7.3 to 0.1) and −21.7 (−27.3 to −16.1) g/L, respectively (
      • Rabaza A.
      • Fraga M.
      • Mendoza A.
      • Giannitti F.
      A meta-analysis of the effects of colostrum heat treatment on colostral viscosity, immunoglobulin G concentration, and the transfer of passive immunity in newborn dairy calves.
      ). Further, in a study by
      • Kryzer A.A.
      • Godden S.M.
      • Schell R.
      Heat-treated (in single aliquot or batch) colostrum outperforms non-heat-treated colostrum in terms of quality and transfer of immunoglobulin G in neonatal Jersey calves.
      , batch heat treatment resulted in a lower total plate count but a higher total coliform count when compared with single bag heat treatment. Notably, neither batch nor bag heat-treatment result in a sterile product and some bacterial species, such as staphylococci and environmental streptococci, appear more tolerant to survive treatment (
      • Elizondo-Salazar J.A.
      • Jayarao B.M.
      • Heinrichs A.J.
      Effect of heat treatment of bovine colostrum on bacterial counts, viscosity, and Immunoglobulin G concentration.
      ,
      • Mann S.
      • Curone G.
      • Chandler T.
      • Moroni P.
      • Cha J.
      • Bhawal R.
      • Zhang S.
      Heat treatment of bovine colostrum: I. Effects on bacterial and somatic cell counts, immunoglobulin, insulin, and IGF-I concentrations, as well as the colostrum proteome.
      ). In a study by
      • Godden S.
      • McMartin S.
      • Feirtag J.
      • Stabel J.
      • Bey R.
      • Goyal S.
      • Metzger L.
      • Fetrow J.
      • Wells S.
      • Chester-Jones H.
      Heat-treatment of bovine colostrum. II: effects of heating duration on pathogen viability and immunoglobulin G.
      , heat treating colostrum at 60°C for 60 min. was successful in eliminating inoculated viable infectious agents Mycoplasma bovis (108 cfu/mL), Listeria monocytogenes (106 cfu/mL), Escherichia coli (106 cfu/mL), and Salmonella enteritidis (106 cfu/mL), but Mycobacterium avium subspecies paratuberculosis, the agent causing Johne's disease, inoculated at 103 cfu/mL was recovered in 1 of 4 batches of colostrum. Further, Staphylococcus aureus and coliforms were not detected after heat treating colostrum 60°C for 60 min. (
      • Elizondo-Salazar J.A.
      • Jayarao B.M.
      • Heinrichs A.J.
      Effect of heat treatment of bovine colostrum on bacterial counts, viscosity, and Immunoglobulin G concentration.
      ). The effectiveness of heat treatment to eliminate viable infectious agents in colostrum depends on the initial pathogen load (
      • Mann S.
      • Curone G.
      • Chandler T.
      • Moroni P.
      • Cha J.
      • Bhawal R.
      • Zhang S.
      Heat treatment of bovine colostrum: I. Effects on bacterial and somatic cell counts, immunoglobulin, insulin, and IGF-I concentrations, as well as the colostrum proteome.
      ). As such, preventing contamination during colostrum harvest and storage should remain a priority regardless of the use of heat-treatment. Additionally, heat-treatment of colostrum alone was not effective at decreasing the risk of Mychobacterium avium subspecies paratuberculosis transmission (
      • Godden S.M.
      • Wells S.
      • Donahue M.
      • Stabel J.
      • Oakes J.M.
      • Sreevatsan S.
      • Fetrow J.
      Effect of feeding heat-treated colostrum on risk for infection with Mycobacterium avium ssp. paratuberculosis, milk production, and longevity in Holstein dairy cows.
      ). Since colostrum only accounts for one potential source of pathogen exposure, measures should be taken to identify and control other routes of disease transmission concurrently.
      Although heat-treatment is an effective strategy to reduce bacterial counts, recent evidence suggests it also alters other colostral components. When inoculating sterile colostrum with fecal E. coli, we observed a greater bacterial growth from 4 to 24 h in heat-treated compared with raw or frozen colostrum (McKane et al., personal communication) suggesting heat treatment decreases the bacteriostatic or bactericidal properties of colostrum. Heat-treatment also increased colostrum viscosity (
      • Rabaza A.
      • Fraga M.
      • Mendoza A.
      • Giannitti F.
      A meta-analysis of the effects of colostrum heat treatment on colostral viscosity, immunoglobulin G concentration, and the transfer of passive immunity in newborn dairy calves.
      ), altered the profile of colostral proteins and metabolites (
      • Tacoma R.
      • Gelsinger S.
      • Lam Y.
      • Scuderi R.
      • Ebenstein D.
      • Heinrichs A.
      • Greenwood S.
      Exploration of the bovine colostrum proteome and effects of heat treatment time on colostrum protein profile.
      ,
      • Mann S.
      • Curone G.
      • Chandler T.
      • Moroni P.
      • Cha J.
      • Bhawal R.
      • Zhang S.
      Heat treatment of bovine colostrum: I. Effects on bacterial and somatic cell counts, immunoglobulin, insulin, and IGF-I concentrations, as well as the colostrum proteome.
      ,
      • Xu W.
      • Mann S.
      • Curone G.
      • Kenéz Á.
      Heat treatment of bovine colostrum: effects on colostrum metabolome and serum metabolome of calves.
      ), as well as reduced or eliminated reactive oxygen and nitrogen species (
      • Mann S.
      • Gandy J.
      • Curone G.
      • Abuelo A.
      The effect of heat treatment on colostral and newborn calf redox status and oxylipid biomarkers.
      ) and active immune components such as colostral leukocytes and reduced activity of the alternative complement pathway (
      • Chandler T.
      • Newman A.
      • Cha J.
      • Sipka A.
      • Mann S.
      Leukocytes, microRNA, and complement activity in raw, heat-treated, and frozen colostrum and their dynamics as colostrum transitions to mature milk in dairy cows.
      ).
      Further, heat-treatment reduced colostral insulin and IGF-1 concentrations (
      • Mann S.
      • Curone G.
      • Chandler T.
      • Moroni P.
      • Cha J.
      • Bhawal R.
      • Zhang S.
      Heat treatment of bovine colostrum: I. Effects on bacterial and somatic cell counts, immunoglobulin, insulin, and IGF-I concentrations, as well as the colostrum proteome.
      ) both of which are important for neonatal gastrointestinal tract development (
      • Ontsouka E.C.
      • Albrecht C.
      • Bruckmaier R.M.
      Invited review: Growth-promoting effects of colostrum in calves based on interaction with intestinal cell surface receptors and receptor-like transporters.
      ,
      • Hammon H.
      • Liermann W.
      • Frieten D.
      • Koch C.
      Importance of colostrum supply and milk feeding intensity on gastrointestinal and systemic development in calves.
      ,
      • Fischer-Tlustos A.J.
      • Cabus A.J.L.
      • Hare K.S.
      • Wood K.
      • Steele M.
      Invited Review: Effects of colostrum management on transfer of passive immunity and the potential role of colostral bioactive components on neonatal calf development and metabolism.
      ) but also has been shown to increase prebiotics such as free oligosaccharides (
      • Fischer A.J.
      • Malmuthuge N.
      • Guan L.L.
      • Steele M.A.
      Short communication: The effect of heat treatment of bovine colostrum on the concentration of oligosaccharides in colostrum and in the intestine of neonatal male Holstein calves.
      ). Despite this, feeding colostrum heat-treated at ≤ 60°C to calves resulted in a 2.5 to 6.6 g/L increase in circulating IgG concentration and a 3.8 to 11.5% increase in apparent efficacy of IgG absorption (AEA;
      • Elizondo-Salazar J.A.
      • Heinrichs A.J.
      Feeding heat-treated colostrum to neonatal dairy heifers: Effects on growth characteristics and blood parameters.
      ,
      • Hesami S.
      • Shahraki A.-D.F.
      • Zakian A.
      • Ghalamkari G.
      The effect of various heat-treatment methods on colostrum quality, health and performance of dairy calves.
      ,
      • Rabaza A.
      • Fraga M.
      • Mendoza A.
      • Giannitti F.
      A meta-analysis of the effects of colostrum heat treatment on colostral viscosity, immunoglobulin G concentration, and the transfer of passive immunity in newborn dairy calves.
      ) as well as a lower risk of preweaning treatment [189 (36.5%) vs. 171 (30.9%);
      • Godden S.
      • Smolenski D.
      • Donahue M.
      • Oakes J.
      • Bey R.
      • Wells S.
      • Sreevatsan S.
      • Stabel J.
      • Fetrow J.
      Heat-treated colostrum and reduced morbidity in preweaned dairy calves: Results of a randomized trial and examination of mechanisms of effectiveness.
      ] . However, data from
      • Gelsinger S.L.
      • Jones C.M.
      • Heinrichs A.J.
      Effect of colostrum heat treatment and bacterial population on immunoglobulin G absorption and health of neonatal calves.
      demonstrated that regardless of heat-treatment, calves fed colostrum with a high bacteria count had a 9.5 to 12.2 g/L lower circulating IgG concentration at 48 h and a 18.7 to 19.9% lower AEA, emphasizing the need to minimize bacterial contamination at time of feeding. In addition to proper sanitation, a combination with heat treatment at 60°C for 60 min or use of an approved colostrum additive (reviewed by
      • Denholm K.
      A review of bovine colostrum preservation techniques.
      ; verify local regulations for approved colostrum additives allowed as a feed additive) should be considered as strategies where needed to secure calf health.

      Storage

      On-farm storage of colostrum is a critical component of a colostrum management system to preserve colostrum composition and IgG concentration as well as to maintain an adequate supply through seasonal declines in yield. Unless fresh colostrum is fed immediately, it should be rapidly cooled before entering storage in the refrigerator (4°C) or freezer (−20°C). Colostrum stored at room temperature had greater bacteria counts by 6 h and 42 times more bacteria by 48 h compared colostrum stored in the refrigerator (
      • Cummins C.
      • Lorenz I.
      • Kennedy E.
      The effect of storage conditions over time on bovine colostral immunoglobulin G concentration, bacteria, and pH.
      ,
      • Cummins C.
      • Berry D.
      • Murphy J.
      • Lorenz I.
      • Kennedy E.
      The effect of colostrum storage conditions on dairy heifer calf serum immunoglobulin G concentration and preweaning health and growth rate.
      ). Although storing colostrum in the refrigerator does not alter IgG concentration, bacterial counts continued to increase over time (
      • Cummins C.
      • Lorenz I.
      • Kennedy E.
      The effect of storage conditions over time on bovine colostral immunoglobulin G concentration, bacteria, and pH.
      ). However, use of potassium sorbate as a colostrum preservative in combination with refrigeration has been an effective strategy to reduce bacterial growth for 96 h compared with raw colostrum in the refrigerator (
      • Stewart S.
      • Godden S.
      • Bey R.
      • Rapnicki P.
      • Fetrow J.
      • Farnsworth R.
      • Scanlon M.
      • Arnold Y.
      • Clow L.
      • Mueller K.
      • Ferrouillet C.
      Preventing bacterial contamination and proliferation during the harvest, storage, and feeding of fresh bovine colostrum.
      ). Because of this, it is recommended that colostrum is stored in the refrigerator for ≤ 2 d or ≤ 4 d when treated with potassium sorbate (
      • Stewart S.
      • Godden S.
      • Bey R.
      • Rapnicki P.
      • Fetrow J.
      • Farnsworth R.
      • Scanlon M.
      • Arnold Y.
      • Clow L.
      • Mueller K.
      • Ferrouillet C.
      Preventing bacterial contamination and proliferation during the harvest, storage, and feeding of fresh bovine colostrum.
      ). For long-term storage, colostrum can be frozen at −20°C (
      • Carlson S.M.A.
      • Muller L.D.
      Compositional and Metabolic Evaluation of Colostrum Preserved by Four Methods during Warm Ambient Temperatures.
      ,
      • Schipper I.
      • Kotta P.
      • Staples G.
      • Fisher G.
      • Erickson G.
      Immunoglobulin-G content in bovine colostrum preserved by freezing, fermentation and chemical preservatives.
      ). Limited data are available to determine the effect of extended storage time on colostral components. In a study by
      • Abd El-Fattah A.M.
      • Abd Rabo F.H.R.
      • El-Dieb S.M.
      • Satar El-Kashef H.A.
      Preservation methods of buffalo and bovine colostrum as a source of bioactive components.
      , concentrations of IgG and IgM in bovine colostrum were not affected when storing colostrum in a freezer for 3 mo, but IgG and IgM decreased 14.6 and 60.5%, respectively when stored for 6 mo. Notably, IgG (30.2 ± 3.0 g/L) and IgM (3.0 ± 0.1 g/L) concentrations in the aforementioned study were lower than that typically observed. Freezing human colostrum preserved concentrations of epidermal growth factor, transforming growth factor (TGF)-β2, tumor necrosis factor (TNF)-α, TNF-receptor I, interleukin (IL)-6, IL-10 for 12 mo; however, IgA, IL-8, and TGF- β1 were only stable when frozen for 6 mo (
      • Ramírez-Santana C.
      • Pérez-Cano F.
      • Audí C.
      • Castell M.
      • Moretones M.
      • López-Sabater M.
      • Castellote C.
      • Franch A.
      Effects of cooling and freezing storage on the stability of bioactive factors in human colostrum.
      ). Notably, ≥ 2 freeze-thaw cycles has been shown to reduce IgG concentration (
      • Morrill K.
      • Robertson K.
      • Spring M.
      • Robinson A.
      • Tyler H.
      Validating a refractometer to evaluate immunoglobulin G concentration in Jersey colostrum and the effect of multiple freeze–thaw cycles on evaluating colostrum quality.
      ) and ≥ 3 freeze-thaw cycles reduced Brix % (
      • Stalker J.M.
      • Gamsjäger L.
      • Pearson J.M.
      • Morck D.W.
      • Windeyer M.C.
      The impact of freezing and multiple freeze-thaw cycles on Brix refractometry estimates of immunoglobulin concentration in beef cattle colostrum.
      ). Therefore, colostrum should be frozen in individual meal portions in a manual defrosting freezer. To avoid raising the internal temperature of the freezer, colostrum can be cooled with ice or cold water before entering the freezer and care should be taken to avoid contact between thawed and frozen containers or bags. Colostrum can be thawed/reheated in a water bath at ≤ 60°C (
      • Balthazar E.
      • Doligez E.
      • Leray O.
      • Le Cozler Y.
      A comparison of thawing methods on IgG1 concentration in colostrum of dairy cows.
      ) without negative effects on serum IgG concentration in the calf (
      • Holloway N.M.
      • Tyler J.W.
      • Lakritz J.
      • Carlson S.L.
      • Holle J.
      Serum immunoglobulin G concentrations in calves fed fresh and frozen colostrum.
      ,
      • Donovan D.C.
      • Reber A.J.
      • Gabbard J.D.
      • Aceves-Avila M.
      • Galland K.L.
      • Holbert K.A.
      • Ely L.O.
      • Hurley D.J.
      Effect of maternal cells transferred with colostrum on cellular responses to pathogen antigens in neonatal calves.
      ). While some authors have not reported reduced concentrations of IgG when thawing colostrum in a microwave (
      • Jones L.R.
      • Taylor A.W.
      • Hines H.
      Characteristics of frozen colostrum thawed in a microwave oven.
      ,
      • Pfeiffer J.
      • Stucke T.
      • Freitag M.
      Effects of different thawing regimes for cow colostrum on the function of colostral immunoglobulin G.
      ),
      • Balthazar E.
      • Doligez E.
      • Leray O.
      • Le Cozler Y.
      A comparison of thawing methods on IgG1 concentration in colostrum of dairy cows.
      observed a 20 to 31% loss of IgG. Heating colostrum in the microwave resulted in coagulation likely from uneven heating as well as reduced volume and crude protein (
      • Jones L.R.
      • Taylor A.W.
      • Hines H.
      Characteristics of frozen colostrum thawed in a microwave oven.
      ). As with heat-treatment, exposing colostrum to temperatures > 60°C when thawing can lead to reductions in IgG concentration.
      Recently, we have shown freezing colostrum short-term increased abundance of microRNA as well as preserved the activity of the alternative complement pathway and bacteriostatic or bactericidal properties compared with raw colostrum (
      • Chandler T.
      • Newman A.
      • Cha J.
      • Sipka A.
      • Mann S.
      Leukocytes, microRNA, and complement activity in raw, heat-treated, and frozen colostrum and their dynamics as colostrum transitions to mature milk in dairy cows.
      ; McKane et al., personal communication). Similar preservation of bacteriostatic or bactericidal properties have been reported with breast milk after freezing although the activity may decrease with extended storage (
      • Ogundele M.O.
      Techniques for the storage of human breast milk: implications for anti-microbial functions and safety of stored milk.
      ,
      • Ogundele M.O.
      Effects of storage on the physicochemical and antibacterial properties of human milk.
      ,
      • Lorico J.L.L.
      • Perez M.L.
      • ng Makati O.
      Effects of storage process on the bacterial growth-inhibiting activity of expressed human breast milk on common neonatal pathogens, Staphylococcus aureus, Escherichia coli and Klebsiella pneumoniae.
      ). Further, freezing colostrum did not alter odd or branch chain fatty acids (
      • Xin H.
      • Xu Y.
      • Chen Y.
      • Chen G.
      • Steele M.A.
      • Guan L.L.
      Odd-chain and branched-chain fatty acid concentrations in bovine colostrum and transition milk and their stability under heating and freezing treatments.
      ), lactoferrin concentrations (
      • Holloway N.M.
      • Lakritz J.
      • Tyler J.W.
      • Carlson S.L.
      Serum lactoferrin concentrations in calves fed fresh or frozen colostrum.
      ), or the antioxidant capacity (
      • Usuga A.
      • Zabala D.A.
      • Medina L.C.
      • Hernandez D.V.
      • Ramirez W.V.
      • Rojano B.A.
      Effect of three storage methods on physical and chemical properties of colostrum from Bos indicus cows.
      ), and was associated with a 1 log reduction in Mycoplasma bovis titer (
      • Gille L.
      • Boyen F.
      • Van Driessche L.
      • Valgaeren B.
      • Haesebrouck F.
      • Deprez P.
      • Pardon B.
      Effect of freezer storage time and thawing method on the recovery of Mycoplasma bovis from bovine colostrum.
      ) but viable maternal leukocytes were eliminated after short-term freezing of colostrum (
      • Chandler T.
      • Newman A.
      • Cha J.
      • Sipka A.
      • Mann S.
      Leukocytes, microRNA, and complement activity in raw, heat-treated, and frozen colostrum and their dynamics as colostrum transitions to mature milk in dairy cows.
      ).

      TRANSITION MILK

      Because calves suckling their dams would naturally experience a gradual decline in nutrient density as well as bioactive concentrations in milk during the first few days of life (
      • Fischer A.
      • Villot C.
      • van Niekerk J.
      • Yohe T.
      • Renaud D.
      • Steele M.
      Invited Review: Nutritional regulation of gut function in dairy calves: From colostrum to weaning.
      ,
      • Tortadès M.
      • Garcia-Fruitós E.
      • Arís A.
      • Terré M.
      Short communication: The biological value of transition milk: analyses of immunoglobulin G, IGF-I and lactoferrin in primiparous and multiparous dairy cows.
      ), feeding transition milk (milkings 2–6) to calves after feeding colostrum has been explored. Feeding transition milk for 1 to 3 d in addition to colostrum resulted in an increased preweaning weight gain (
      • Van Soest B.
      • Cullens F.
      • VandeHaar M.
      • Nielsen M.W.
      Effects of transition milk and milk replacer supplemented with colostrum replacer on growth and health of dairy calves.
      ) and lower odds of receiving an elevated ear/eye or nasal score (
      • Conneely M.
      • Berry D.P.
      • Murphy J.P.
      • Lorenz I.
      • Doherty M.L.
      • Kennedy E.
      Effect of feeding colostrum at different volumes and subsequent number of transition milk feeds on the serum immunoglobulin G concentration and health status of dairy calves.
      ). Further, calves fed a 1:1 colostrum:whole milk mixture for 3 d after a colostrum meal had increased intestinal surface area as well as increased villi height compared with calves fed whole milk (
      • Pyo J.
      • Hare K.
      • Pletts S.
      • Inabu Y.
      • Haines D.
      • Sugino T.
      • Guan L.
      • Steele M.
      Feeding colostrum or a 1: 1 colostrum: milk mixture for 3 days postnatal increases small intestinal development and minimally influences plasma glucagon-like peptide-2 and serum insulin-like growth factor-1 concentrations in Holstein bull calves.
      ). Despite its benefit to the calf, feeding transition milk on a commercial dairy farm presents an added management challenge because of the additional equipment and labor required to harvest and feed transition milk to a select group of calves. Research seeking to determine the optimum duration to feed transition milk as well as identifying the short- and long-term benefits to health, growth, development, and future productivity and compare these outcomes when calves are fed maternal transition milk to protocols feeding whole milk and to those using colostrum supplements to replace transition milk will be instrumental in adoption of transition milk programs on commercial dairy farms. Transition milk can be harvested from cows 2 to 6 milkings after calving and can be fed fresh, heat-treated, or stored as described above for colostrum. Further, transition milk can be pooled from multiple cows; however, measures should be taken to lower the risk of disease transmission when pooling transition milk in the same fashion as when pooling colostrum from multiple dams. Alternatively, mixing colostrum supplements with milk has been used as a substitution for harvesting transition milk (
      • Berge A.C.B.
      • Besser T.E.
      • Moore D.A.
      • Sischo W.M.
      Evaluation of the effects of oral colostrum supplementation during the first fourteen days on the health and performance of preweaned calves.
      ).

      FUTURE OPPORTUNITIES

      Despite the variables summarized herein, the proportion of variance in colostrum production we have been able to explain remains small, in part, due to the incomplete knowledge of the physiological mechanisms of colostrum formation (Figure 1). Because of the importance of and traditional focus on IgG alone, transfer of IgG into the mammary gland has historically defined colostrogenesis and is believed to begin 3 to 5 weeks before calving when IgG concentrations in the mammary gland exceed the concentration in maternal circulation (
      • Brandon M.
      • Watson D.
      • Lascelles A.
      The mechanism of transfer of immunoglobulin into mammary secretion of cows.
      ,
      • Baumrucker C.R.
      • Bruckmaier R.M.
      Colostrogenesis: IgG1 transcytosis mechanisms.
      ). During the days to weeks leading up to calving, nutritional and non-nutritional components accumulate in the mammary secretion (
      • Hurley W.
      Mammary function during the nonlactating period: enzyme, lactose, protein concentrations, and pH of mammary secretions.
      ,
      • Bitman J.
      • Wood D.
      • Capuco A.V.
      Changes in the lipid composition of the secretions of the bovine mammary gland during the dry period.
      ,
      • Guy M.A.
      • McFadden T.B.
      • Cockrell D.C.
      • Besser T.E.
      Regulation of colostrum formation in beef and dairy cows.
      ,
      • Baumrucker C.R.
      • Gross J.J.
      • Bruckmaier R.M.
      The importance of colostrum in maternal care and its formation in mammalian species.
      ,
      • Hare K.
      Relationships between maternal nutrition, colostrogenesis, and neonatal physilogical and metabolic development in cattle.
      ) and numerous hormones (estradiol, progesterone, prolactin, cortisol, leptin, placental lactogen, etc.) have been discussed as having a potential role in colostrum formation as well as the initiation of lactogenesis II (reviewed by
      • Barrington G.
      • McFadden T.
      • Huyler M.
      • Besser T.
      Regulation of colostrogenesis in cattle.
      ,
      • Baumrucker C.R.
      • Macrina A.L.
      • Bruckmaier R.M.
      Colostrogenesis: Role and Mechanism of the Bovine Fc Receptor of the Neonate (FcRn).
      , Bigler et al., 2023). However, the timing and complete cascade of signals that influence these biological mechanisms, as well as active and passive transfer of constituents remain unknown. Understanding these signals as well as the onset of lactogenesis II are particularly important as the capacity, rate, and time of which colostral components enter or are synthesized in the mammary gland could influence the osmotic gradient and as such affect yield as well as the concentration of colostral components.
      Figure thumbnail gr1
      Figure 1Summary of cow, colostrum harvest, and post-harvest variables identified as focus of future colostrum investigations. Figure was created with BioRender.
      The ability to improve colostrum production through prepartum nutrition and management as well as genetic selection remains a plausible and achievable goal. However, because of the high variability observed in colostrum production and limited available data, future research is needed to discover new interventions that directly influence colostrum yield and composition and increase the external validity of existing findings. Further, inclusion of colostrum outcomes and consistent reporting among future transition cow investigations will facilitate subsequent meta-analyses of these typically smaller studies. The mechanisms of how nutritional or management interventions affect colostrum production remains unclear. Attention to the interactions with maternal metabolism and endocrine signals is necessary to understand these regulatory mechanisms.
      Lastly, data on the role of harvest and post-harvest management on colostral components as well the effect on health, growth, and future productivity of calves fed colostrum are needed. Research investigating whether harvest procedures for colostrum and transition milk should differ from the procedures used to harvest mature milk are limited at this time. Further, recording colostrum yield and Brix % readings into dairy management software on commercial dairy farms will enable high-powered and externally valid observational data analysis to identify risk factors and genomic trends. Recent evidence suggests heat-treatment, bacterial contamination, refrigeration, and freezing affect colostral components or the absorption of IgG in the calf (
      • Ramírez-Santana C.
      • Pérez-Cano F.
      • Audí C.
      • Castell M.
      • Moretones M.
      • López-Sabater M.
      • Castellote C.
      • Franch A.
      Effects of cooling and freezing storage on the stability of bioactive factors in human colostrum.
      ,
      • Tacoma R.
      • Gelsinger S.
      • Lam Y.
      • Scuderi R.
      • Ebenstein D.
      • Heinrichs A.
      • Greenwood S.
      Exploration of the bovine colostrum proteome and effects of heat treatment time on colostrum protein profile.
      ,
      • Fischer A.J.
      • Malmuthuge N.
      • Guan L.L.
      • Steele M.A.
      Short communication: The effect of heat treatment of bovine colostrum on the concentration of oligosaccharides in colostrum and in the intestine of neonatal male Holstein calves.
      ,
      • Mann S.
      • Curone G.
      • Chandler T.
      • Moroni P.
      • Cha J.
      • Bhawal R.
      • Zhang S.
      Heat treatment of bovine colostrum: I. Effects on bacterial and somatic cell counts, immunoglobulin, insulin, and IGF-I concentrations, as well as the colostrum proteome.
      ,
      • Chandler T.
      • Newman A.
      • Cha J.
      • Sipka A.
      • Mann S.
      Leukocytes, microRNA, and complement activity in raw, heat-treated, and frozen colostrum and their dynamics as colostrum transitions to mature milk in dairy cows.
      ,
      • Mann S.
      • Gandy J.
      • Curone G.
      • Abuelo A.
      The effect of heat treatment on colostral and newborn calf redox status and oxylipid biomarkers.
      ,
      • Stalker J.M.
      • Gamsjäger L.
      • Pearson J.M.
      • Morck D.W.
      • Windeyer M.C.
      The impact of freezing and multiple freeze-thaw cycles on Brix refractometry estimates of immunoglobulin concentration in beef cattle colostrum.
      ). Focusing on the nutritional and developmental role of other colostral components and how current post-harvest management affects these components might redirect best-practice guidelines and redefine colostrum quality. Moreover, the use of colostrum as a therapeutic as well as research into the success of treatment protocols using colostrum are needed (
      • Carter H.S.
      • Renaud D.L.
      • Steele M.A.
      • Fischer-Tlustos A.J.
      • Costa J.H.
      A narrative review on the unexplored potential of colostrum as a preventative treatment and therapy for diarrhea in neonatal dairy calves.
      ,
      • Carter H.S.M.
      • Steele M.A.
      • Costa J.H.C.
      • Renaud D.L.
      Evaluating the effectiveness of colostrum as a therapy for diarrhea in preweaned calves.
      ). Lastly, the effect of feeding transition milk as well as practical strategies to harvest and feed transition milk on a commercial farm warrants further investigation.

      CONCLUSIONS

      Colostrum yield and composition exhibit individual, herd, and seasonal variability. Although multiple animal and environmental variables have been linked with colostrum production, researchers have been mostly unsuccessful in explaining this variability. Prepartum nutrition and management as well as the interaction with maternal metabolism appear to affect colostrum production. However, suitable on-farm strategies to improve colostrum production remain limited, partially because of our incomplete knowledge on the regulatory mechanisms of colostrum formation. Storage of colostrum remains an effective approach to overcome periods of low colostrum supply. Post-harvest colostrum management should limit bacterial contamination and future studies need to quantify the effect on colostral components and ultimately on calf health.

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