Abstract

Research on effects of water-soluble vitamins when fed to dairy cows and field supplementation of some water soluble vitamins has increased markedly in the past few years. In research studies, biotin supplementation (20 mg/day) has consistently improved hoof health of cows and often increased milk production. Supplementation of niacin, although common, has little effect on milk production unless the supplementation rate is 12 g/day, and then the response is often not profitable. Adding rumen-protected choline (50 g/day) often increases milk production in early lactation, and the average response is profitable. Research on folic acid, vitamin B-12, and vitamin C for dairy cows is continuing, but at the present time inadequate data are available to recommend routine supplementation.

Introduction

Vitamins are organic compounds needed in minute amounts that are essential for life. A vitamin must be in the diet (dietary essential) or be synthesized by microorganisms in the digestive system and then absorbed by the host animal. Currently there are 14 recognized vitamins of which four are fat-soluble and 10 are water-soluble, but not all animals require all 14 vitamins (Table 1). When an animal absorbs an inadequate quantity of a particular vitamin, various responses are observed depending on the vitamin and the degree and duration of deficiency. The most severe situation (seldom observed in U.S. dairy cows) is a clinical deficiency. For example, rickets results from a clinical deficiency of vitamin D. Marginal deficiencies of vitamins usually have more subtle and less defined signs. Unthriftiness; reduced growth rate, milk production, or fertility; and increased prevalence of infectious diseases can be observed when animals absorb inadequate amounts of vitamins.

It is not known definitively whether cows have an absolute dietary requirement for any of the water-soluble vitamins. The liver and kidney of the cow can synthesize vitamin C, and ruminal and intestinal bacteria synthesize most, if not all, of the B vitamins. The concentrations of many B vitamins are relatively high in many common feeds; therefore, in the vast majority of situations, cows do not need to consume any supplemental water-soluble vitamins to prevent clinical deficiency. In a survey (Kellogg et al., 2001) of the highest producing dairy herds in the United States (data collected in 2000), niacin was the only water-soluble vitamin fed to a substantial number of herds (43% of the herds reported that at least one group of cows was fed niacin). Choline and biotin were also fed, but they were used by less than 4% of the surveyed herds. Even though clinical deficiencies of water-soluble vitamins are extremely rare in dairy cows, research and field interest in water-soluble vitamins has increased markedly in the past few years.

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Why the Increased Interest in Water-Soluble Vitamins?

The predominant function of the B vitamins is to act as co-factors for enzymes that are involved in amino acid, energy, fatty acid, and nucleic acid metabolism (Table 1). Many of these enzymes are involved directly in the production of milk and milk components. Therefore, as milk production increases, the need for these enzymes (and the associated co-factors) increase. In the past 15 years, average milk yield per cow has increased from about 14,500 lb per year to almost 19,500 lb, and herds (not individual cows) that average 28,000 lb or more per cow are not uncommon. Assuming average milk composition (for a Holstein) and assuming composition has not changed over time, the average cow in 2005 must synthesize approximately 0.4 lb more milk fatty acids (assuming 50% of milk fatty acids comes from the diet), 0.6 lb more milk protein, and 0.9 lb more lactose each day than the average cow in 1990. During that same 15-year period, average dry matter intake has increased from about 44 lb to about 50 lb/day. In other words, the yield of milk and milk components has increased about 33%, but dry matter intake has increased only about 15%. Because most B vitamins are not supplemented, supply to the cow would mostly be a function of intake, whereas their need would be a function of milk production. The potential imbalance between supply and need in today’s high-producing cow increases the likelihood that responses will be observed when B vitamins are supplemented. The interest in B vitamins has increased because cows have changed.

B Vitamin Supply

As with all nutrients, a response to supplemental B vitamins will only be observed if: 1) supplementation actually increases the supply of vitamin to the tissues that require it, and 2) the nutrient is first limiting. Vitamin supply is defined as the amount (micrograms or milligrams) of a vitamin that is absorbed from the digestive system each day. Supply is a function of the amount of the vitamin consumed (vitamin concentration times dry matter intake), ruminal synthesis and degradation of the vitamin, and its bioavailability (i.e., its ability to be absorbed, mainly by the small intestine).

Dietary Concentrations

Because of the difficulty in measuring many of the B vitamins, we have very limited data on their concentrations in common feedstuffs. Inadequate information is available to discuss differences in concentrations of B vitamins among feedstuffs, but data for some feeds are available (Schwab et al., 2006). Ranges in reported concentrations of various B vitamins in diets fed to lactating cows are in Table 2. Most of the data represent seven relatively diverse diets from three experiments (Santschi et al., 2005a; Santschi et al., 2005b; Schwab et al., 2006), but it is important to note that all the analyses were conducted in a single laboratory. Considering the analytical and sampling error usually observed when trace nutrients are measured, concentrations of most of the B vitamins were relatively consistent across the diverse diets with the clear exception of niacin. The concentration of dietary niacin was mostly a function of the amount of soyhulls included in the diet. In the four diets that contained little or no soyhulls, niacin concentrations were <30 mg/kg, and in the three diets that contained appreciable concentrations of soyhulls, niacin concentrations were >60 mg/kg. Additional data are needed to confirm whether soyhulls typically contain such high concentrations of niacin. The biotin concentration of different diets within an analytical method did not vary greatly, but method of analysis had a substantial effect (Table 2). Biotin concentrations in three studies that used one analytical procedure averaged about 7 mg/kg, and in three other studies using a different procedure. it was almost 20 times lower (about 0.4 mg/kg). At the current time, we do not know which method is accurate.

Ruminal Metabolism

The flow of B vitamins measured at the duodenum can be substantially different from intake (Zinn et al., 1987; Santschi et al., 2005a; Schwab et al., 2006). The difference between intake and duodenal flow is called net synthesis because it reflects both ruminal degradation and synthesis. For most B vitamins, flow out of the rumen exceeds intake, indicating net synthesis of the vitamin (Table 3). With the exception of biotin and vitamin B-6, ruminal synthesis appears to provide the majority of the B vitamins that reach the small intestine (Table 3). Both studies that used dairy cows (Santschi et al., 2005a; Schwab et al., 2006) reported no net synthesis of biotin or that ruminal degradation was slightly higher than synthesis (i.e., flow to the duodenum was statistically lower than biotin intake). Those two studies also were among those that reported very high concentrations of biotin in the diet (Table 2). If the concentration of biotin in the diet was overestimated, net synthesis would be underestimated. Net ruminal synthesis of biotin in beef cattle (Miller et al., 1986; Zinn et al., 1987) and in in vitro ruminal systems (Abel et al., 2001) was positive. In addition, the development of clinical signs of biotin deficiency was prevented when chicks were fed ruminal contents but not when fed the same diet fed to the cow from which the ruminal contents were obtained (McElroy and Jukes, 1940). Additional research is needed to clarify the question regarding biotin synthesis in the rumen.

Only one study that used modern analytical techniques has evaluated how dietary factors influenced duodenal flow of B vitamins (Schwab et al., 2006). In that study, diets had either 40 or 60% forage (50:50 mix of corn silage and hay) with low (approximately 6.5%) or moderate (approximately 20%) starch concentrations. Starch concentration was varied by replacing dry ground corn with soyhulls. Cows fed the low forage diets consumed about 5 lb/day more (P < 0.01) dry matter than cows fed the high forage diet, and duodenal flow of thiamin, niacin, B-6, folic acid, and B-12 was also higher. Much of the increased flow of those vitamins was a direct result of increased intake, but apparent ruminal synthesis of folic acid and B-12 was also increased when low forage diets were fed. Cows fed the moderate starch diets consumed more dry matter than cows fed the low starch diets, and duodenal flows of B-6, biotin, and folic acid were also higher. Apparent ruminal synthesis of niacin was more than doubled when the moderate starch diets were fed, but because the low starch diets contained much higher concentrations of niacin, duodenal flow was not affected. Apparent ruminal synthesis of folic acid and B-6 was increased with moderate starch diets, but synthesis of B-12 was reduced. Overall, it appears that ruminal synthesis of most B vitamins is related to microbial fermentation in the rumen. Diets that have a higher concentration of rumen fermentable matter promote increased synthesis of many of the B vitamins but may reduce synthesis of B-12.

Apparent ruminal synthesis of B vitamins equals: [synthesis of the vitamins by ruminal microorganisms – (degradation of the vitamins by ruminal microorganisms + ruminal and omasal absorption of the vitamin)]. To measure ruminal disappearance of B vitamins, diets with and without supplemental vitamins are fed, and duodenal flows of the vitamins are compared. If supplementation resulted in no increase in duodenal flow of that vitamin, apparent disappearance equals 100%. Degradation of B vitamins contained in feedstuffs may or may not be the same as disappearance of supplemental B vitamins. Santschi et al. (2005a) measured apparent ruminal disappearance of supplemental B vitamins in dairy cows. Approximately 100% of supplemental riboflavin, niacin, and folic acid disappeared in the rumen. Approximately two-thirds of the supplemental thiamin and B-12 and 40 to 45% of the supplemental B-6 and biotin (the variation in biotin disappearance was extremely high) disappeared. Because apparent ruminal disappearance is caused by both microbial degradation and absorption, high disappearance values do not necessarily mean that responses to supplementation are unlikely. The rumen and omasum do not appear to be major absorptive sites for most B vitamins, but high supplementation rates still might increase systemic concentrations of some vitamins. In addition, some rumen microorganisms require B vitamins; therefore, ruminal effects can occur even if a substantial amount of the supplemental vitamin disappears in the rumen.

Intestinal Absorption

Apparent intestinal absorption is calculated by subtracting the flow of a vitamin at the ileum from flow measured at the duodenum. Bacteria can inhabit the terminal portion of the small intestine (ileum); therefore, apparent intestinal absorption measured in this way would underestimate true absorption if B vitamins are synthesized by those bacteria. In addition, some vitamins that are absorbed are secreted in bile which would result in lower apparent intestinal absorption. Only one study is available that used dairy cows and modern analytical techniques (Santschi et al., 2005a). The researchers measured apparent intestinal absorption of several B vitamins from supplemented and unsupplemented diets. Overall, few differences were observed between supplemented and unsupplemented treatments, suggesting the absorption of basal and supplemental vitamins was similar. Apparent intestinal absorption of thiamin, niacin, and B-6 averaged 70 to 85%; for riboflavin and biotin, it averaged about 35%, and about 13% for B-12. Apparent intestinal absorption of folic acid was negligible probably because of bilary secretion.

Clinical and Production Responses to B Vitamins

Niacin

No requirement for dairy cows has been established for niacin, but niacin is involved in most energy-yielding pathways and for amino acid and fatty acid synthesis and therefore is important for milk production. Niacin has been evaluated for possible prophylactic and therapeutic effects on ketosis and fatty liver syndrome. Although a few studies reported that niacin supplementation during the periparturient period (usually 6 to 12 g/day) reduced blood ketones and plasma nonesterified fatty acids (NEFA), the vast majority of studies (see page 171 of NRC, 2001 for listing of the studies) showed no effect (a few actually found increased ketones and NEFA with niacin supplementation). In a recent study published only in abstract form (French, 2004), Jersey cows were fed 48 g of nicotinic acid/day from 30 d prepartum until calving. The day before calving, cows fed supplemental niacin had greater dry matter intakes (22.0 vs. 14.7 lb) and lower plasma NEFA (491 vs. 1244 µmol/L).

Several summaries of production studies evaluating niacin supplementation have been published (Drackley, 1992; Erdman, 1992; Girard, 1998; NRC, 2001; Schwab et al., 2005). The recent summary by Schwab et al. (2005) was conducted using a new statistical method and is probably the best current summary. They concluded that supplementing 6 g/d of niacin (commonly used supplementation rate) had no effect on milk production or milk composition. At 12 g/d of supplemental niacin, 3.5% fat-corrected milk increased about 1 lb/d, fat yield was increased 26 g/d, and milk protein yield was increased 17 g/d. Based on the current cost of niacin, this response would often not be profitable. The likelihood of a profitable response can be increased by targeting specific animals. Positive responses appear more likely in early lactation, high-producing cows, and responses are almost never observed in mid- and late lactation cows (Girard, 1998). Supplemental niacin often had negative effects when fed with diets that contained supplemental fat (Drackley, 1992). Possible reasons for the limited response to supplemental niacin include: 1) basal diets provide adequate niacin to the intestine (Tables 2 and 3), or 2) supplementation at 6 to 12 g/d does not increase flow of niacin because of extensive ruminal metabolism. Increasing flow of niacin to the duodenum could be accomplished by feeding rumen-protected niacin or perhaps by greatly increasing the supplementation rate. A rumen-protected form of niacin is available, but published data evaluating the product with dairy cows are not available. Additional research is needed to study production and other responses to higher supplementation rates.

Biotin

A dietary biotin requirement has not been established for dairy cows. Six clinical trials have been published that examined the effect of supplemental biotin on hoof horn lesions and lameness in dairy cows (reviewed by Weiss, 2005). Although the response variable varied among experiments, all studies reported reduced prevalence of specific lesions or clinical lameness when biotin was supplemented. The supplementation rate was 20 mg/d in most studies, but one study with beef cows fed only 10 mg/d and reported a positive response, and all studies involved long-term (months) biotin supplementation. Biotin supplementation usually reduces hoof lesions in two to three months, but six months of supplementation may be required to reduce clinical lameness. The mechanisms by which biotin affects foot health are not well understood. Increased keratin synthesis by keratinocytes from the hoof might be a possible mechanism by which biotin improves foot health. Keratinocytes are cells responsible for the synthesis of proteins known as keratins, and keratin synthesis is a main determinant of hoof integrity. Keratin synthesis by human skin keratinocytes was increased when cultured with supra-physiological concentrations of biotin (Fritsche et al., 1991). Increased fatty acid synthesis via increased activity of acetyl-CoA carboxylase might be another mechanism by which biotin improves foot health. The keratinocytes are embedded in a lipid-rich extracellular matrix composed of cholesterol, fatty acids, and ceramides. Higuchi et al. (2004) reported that biotin supplementation decreased the concentration of water and increased the concentration of lipids in the sole of dairy cows.

Milk yield responses to supplemental biotin are less consistent than hoof responses, but the majority of studies reported increased production (Table 4). Low-producing cows and/or cows in late lactation are unlikely to increase milk yield when biotin is supplemented. A recent 14-day study from our laboratory (Ferreira, 2006) found that biotin increased milk yield when supplemented to high-producing dairy cows (control cows average production = 89 lb/day and 13656 days in milk), but not when supplemented to low-producing cows (average production for control cows = 52 lb/day and 26753 days in milk). The lack of a production response by low-producing cows in that study agrees with data from Australia (Fitzgerald et al., 2000). The reason cows in the Rosendo et al. (2004) experiment did not respond is not known (milk production of control cows averaged 79 lb/day). Across all studies, the median increase in milk yield was 2 to 3 lb/day. Whereas months of supplementation are required to observe improved hoof health, the milk yield response appears very rapidly (Figure 1). The mechanism by which biotin supplementation increases milk yield is still not known, but supplemental biotin can increase the activity of one gluconeogenic enzyme in the liver of dairy cows (Ferreira, 2006).

Folic Acid and Vitamin B-12

A substantial amount of research has been conducted in Canada on folic acid and B-12 nutrition of dairy cows (Girard and Matte, 1998; Girard and Matte, 1999; Girard et al., 2005; Girard and Matte, 2005). Vitamin B-12 is essential for folic acid to work properly, and therefore these two vitamins must be considered together. Both vitamins are involved in methionine metabolism. Vitamin B-12 can be synthesized by rumen bacteria if adequate cobalt is in the diet (NRC requirement is 0.11 mg/kg of diet DM, but newer research suggests that 0.2 to 0.3 mg/kg may be better). The effect of folic acid supplementation (typical rates are between about 2 and 3 g/day) on milk production has been variable. In one study, milk production of multiparous cows was increased by 4 to 6 lb/d when folic acid was supplemented, but no effect was observed with first lactation cows. In other experiments, folic acid has not affected milk production. One reason for the variable responses may be that vitamin B-12 status was limiting. If cows are limited in B-12, they are unlikely to respond to folic acid supplementation. Interactions between methionine supply, folic acid, and B-12 are likely. Both vitamin B-12 and folic acid are expensive, and we still do not understand all the factors that influence responses to supplementation. Additional research is needed before routine supplementation of these vitamins is recommended.

Other B Vitamins

Research is extremely limited on the effects of supplementing B vitamins (other than biotin and niacin) to dairy cows. In a study (Majee et al., 2003) in which a mixture of B vitamins (biotin, folic acid, niacin, pantothenic acid, B-6, riboflavin, thiamin, and B-12) was fed, milk production was increased compared with the control but was not different from a treatment in which only biotin was supplemented. When the amount of supplemental B vitamins was doubled, intake and milk production was similar to control cows (i.e., lower than the 1-X supplementation treatment). Shaver and Bal (2000) examined the effects of supplemental thiamin on milk production. In one experiment, yield of milk, milk fat, and milk protein increased when cows were fed 150 mg of thiamin per day. In two other experiments, cows fed thiamin at 300 mg/day had similar milk yields as control cows. Overall, the available data do not support routine supplementation of “other” B vitamins. However, as productivity of cows continues to increase and as new experiments are conducted, this conclusion may change.

Choline

Choline does not fit the definition of a vitamin. It is required in gram quantities (not milligram or microgram quantities), and it is synthesized by the cow. Very little, if any, dietary choline (with the exception of rumen-protected supplements) is absorbed from the gut because it is degraded in the rumen. At the 2002 Tri-State Conference, Donkin (2002) summarized previous data on milk yield responses (10 comparisons) to supplemental choline. Six comparisons (60%) reported a statistical increase in milk production. Two additional studies have since been published, and one paper (Janovick Guretzky et al., 2006) reported no response while the other (Piepenbrink and Overton, 2003) reported increased milk production. Across all 12 comparisons, all but one reported a numerical increase in milk production, and the median increase to choline supplementation was about 5 lb/day. Supplemental choline during the transition period may reduce liver fat, but results have not been consistent. Because choline can be synthesized from methionine, diets that provide marginal amounts of metabolizable methionine may be more likely to respond to choline supplementation. Choline must be rumen-protected to be effective.

Vitamin C

Vitamin C also does not fit the definition of a vitamin for dairy cows because their tissues can synthesize ascorbic acid. Vitamin C is probably the most important water soluble antioxidant in mammals. Most forms of vitamin C are extensively degraded in the rumen (Macleod et al., 1999); therefore, the cow must rely on tissue synthesis of vitamin C. The concentration of ascorbic acid is high in neutrophils and increases as much as 30-fold when the neutrophil is stimulated by the presence of bacteria (Wang et al., 1997). Santos et al. (2001) reported that plasma ascorbic acid concentrations in dairy cows were not correlated with SCC. However, the range in SCC was limited (67,000 to 158,000/ml), and cows were only sampled once. Another experiment evaluated the therapeutic use of ascorbic acid following intramammary challenge with endotoxin (Chaiyotwittayakun et al., 2002). One quarter from each cow was infused with endotoxin, and the ascorbic acid was injected IV at 3 and 5 hours post challenge (25 g/dose). Vitamin C therapy had only limited effects on clinical signs. Because of the way vitamin C works, an endotoxin challenge may not be a very good model to evaluate effects of vitamin C on mastitis. Studies in which mammary glands were either experimentally or naturally infected with bacteria clearly show a relationship between plasma vitamin C concentrations and infection. Cows with a mammary gland infection had lower concentrations of vitamin C in plasma than did healthy cows (Weiss et al., 2004; Kleczkowski et al., 2005; Ranjan et al., 2005). In addition, we (Weiss et al., 2004) observed significant correlations between vitamin C concentrations in plasma and milk and clinical signs of mastitis caused by E. coli. Greater decreases in vitamin C concentrations were related to longer duration of clinical mastitis and greater decreases in milk production. Data from these experiments do not mean that increasing vitamin C status of cows will reduce the prevalence or severity of mastitis. We do not know whether lower vitamin C status allowed cows to become infected or whether the infection depleted body vitamin C.

Conclusions and Recommendations

1. Supplemental biotin provided at about 20 mg/day has consistently improved hoof health and has increased milk production in several, but not all, studies. For improvements in hoof health, biotin must be fed for several months (including the dry period), but increased production will happen within a very short period. Feeding 20 mg/day of biotin to lactating and dry cows is recommended because of its effects on foot health.
2. Rumen-protected choline fed at 50 g/day (actual product, not choline) has resulted in increased milk production in most studies and reduced liver fat in some studies. The cost of supplementation is substantial, but the median response to supplementation was about 5 lb of milk/day. A response in milk production is most likely in early lactation (up to about 60 days in milk); to maximize the likelihood of a profitable return on investment, supplementation should be limited to early lactation cows.
3. A milk production response to niacin supplementation at 6 g/day is unlikely, but supplementation at 12 g/d can increase milk production by about 1 lb (likely not a profitable return on investment). A positive return on investment is more likely when supplementation is limited to early lactation cows. The use of supplemental niacin in herds that feed a single diet to all cows is unlikely to have a positive return on investment.
4. At this time, insufficient data are available to recommend supplementation of other B vitamins and vitamin C to dairy cows.

References

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Tables/Figures

Figure 1. Milk yield response when cows in mid- (136 days in milk) or late (267 days in milk) lactation were supplemented with 20 mg/day of biotin. Dashed lines represent control cows, and solid lines represent supplemented cows. Arrows designate when supplementation started and ended.

Author Information

William P. Weiss and Gonzalo Ferreira
Ohio Agricultural Research and Development Center
The Ohio State University
Wooster, OH 44691
(330) 263-3622, Fax (330) 263-3949
weiss.6@osu.edu