The importance of milk as a source of
vitamin B12 for human nutrition
J. J. Matte,* M. Britten,† and C. L. Girard*
*Agriculture and Agri-Food Canada, Dairy and Swine Research and Development Centre, Sherbrooke, QC, Canada
†Agriculture and Agri-Food Canada, Food Research and Development Centre, St-Hyacinthe, QC, Canada
Implications
• A
 mong animal products, those from ruminants are particularly
rich in vitamin B12, which is naturally synthesized by the ruminal
microflora and transferred to milk.
 oncentrations of vitamin B12 in milk vary considerably and are
• C
affected by diet.
• D
 airy products retain, in general, a major part of the vitamin B12
naturally present in milk, some processing conditions may even
add to the basal level by production of vitamin B12 from propionic
bacterium in Swiss-type cheeses.
• Intestinal bioavailability of vitamin B12 from milk, regardless of
the technological process (raw, pasteurized, or microfiltered) is
greater than the synthetic form used in supplements.
Key words: animal model, bioavailability, cobalamin, cow, milk pro-
cessing
Vitamin B12 in Human Nutrition:
Milk is a Major Source
Among B vitamins, vitamin B12 occupies a very special niche. This
vitamin is produced only by bacteria and archaebacteria if the cobalt
supply is adequate. As opposed to other B vitamins, it is neither synthe-
sized nor used by fungi and plants (Martens et al., 2002). Therefore, in
human diets, the sole natural source of vitamin B12 comes from animal
products. Among animal products, those from ruminants are particularly
rich in vitamin B12, which is naturally synthesized by the ruminal micro-
flora using cobalt as an essential precursor. After intestinal absorption in
the ileum, it is stored in liver and muscles (meat) of the host or secreted
in milk (Combs, 2012).
In humans, vitamin B12 deficiency affects cell division and may lead
to anemia and neuropathy (Combs, 2012). In the presence of vitamin B12
deficiency, increasing folic acid supply cures anemia but not neurologi-
cal symptoms. In fact, by masking the hematological symptoms, folic
acid could delay diagnosis of vitamin B12 deficiency until neurological
damages are irreversible. Consequently, over the last decade, since folic
acid fortification of flour became mandatory in many Western countries,
including Canada, there has been a renewed interest for evaluation of vi-
© Matte, Britten, and Girard.
doi:10.2527/af.2014-0012
32 Animal Frontiers
tamin B12 status in human populations, especially pregnant women and
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the elderly (Selhub et al., 2007, 2009; Selhub and Paul, 2011). In fact, a
Canadian study (Ray et al., 2007) and a recent meta-analysis (Wang et
al., 2012) reported that maternal dietary vitamin B12 would be the major
nutrient related to neural tube defect risks in early pregnancy within
populations receiving supplements of folic acid (periconceptional tab-
lets or fortified foods). As much as 34% of all neural tube defect cases
in Canada might be due to low maternal vitamin B12 status (Ray et al.,
2007). Moreover, especially in the elderly population, it seems that low
vitamin B12 status, even if still within the normal range, is associated
with neurodegenerative disease and cognitive impairment (De Lau et al.,
2009; Moore et al., 2012; Morris et al., 2012). Furthermore, cognitive
decline is accelerated in individuals combining low vitamin B12 and high
folate status (Morris et al., 2012).
Vitamin B12 status is correlated with vitamin B12 intake in humans
(Tucker et al., 2000; Vogiatzoglou et al., 2009; Bor et al., 2010). Veg-
etarians have lower vitamin B12 status than omnivores (Miller et al.,
1991; Bor et al., 2010; Obersby et al., 2013). However, dietary sources
of the vitamin also seem to matter. For example, vitamin B12 status of
vegetarians was positively correlated with their intake of dairy products,
especially milk, but was not correlated with egg or seafood consumption
(Miller et al., 1991). Among adults not using vitamin supplements, the
relationship between plasma concentration of the vitamin and its intake
from dairy products is similar to the relation observed with intake of
cereals fortified with vitamin B12. However, the relationship with meat,
poultry, or fish intakes is weaker (Tucker et al., 2000). A Norwegian
study showed that plasma concentrations of vitamin B12 increase with
the amounts of vitamin B12 provided by dairy products or fish but not
with those provided by eggs or meat (Vogiatzoglou et al., 2009). More-
over, for a similar intake, plasma concentrations of vitamin B12 were
greater when the vitamin was supplied by dairy products compared with
fish, suggesting that bioavailability of the vitamin from dairy products
is greater than for other sources (Vogiatzoglou et al., 2009). Globally,
these dietary surveys seem to indicate that vitamin B12 supplied by dairy
products is more available than from other natural sources, although in
these studies, intake data were obtained by a food-frequency question-
naire. In terms of provision of vitamin B12, one glass (250 mL) of milk
provides more than 1 mg of vitamin B12 (USDA, 2011). According to the
Canadian Food Inspection Agency (2010), cow milk could be labeled as
an “excellent source” of vitamin B12 because one glass of milk provides
nearly 50% of the recommended daily allowance (RDA; Health Canada,
2006) for adults and children over 13 years of age (2.4 mg/day).
 Vitamin B12 in Milk
Influence of the dairy cow and its diet
In 1966, Miller et al. (1966) reported that concentrations of vitamin B12
in milk were highly variable and were affected by cow breed, season, co-
balt supply, and feeding regimens. These authors observed that inclusion
of oat silage increased milk concentrations of vitamin B12 as compared
with corn silage, but details on the studied feeding regimens or intake are
scarce. More recently, in a study comparing four production systems in

With the permission of M. Duplessis

France, mainly characterized by their forage system (grassland or corn
silage) and altitude (lowland or mountain), milk concentrations of vita-
min B12 appeared to be related to the composition of the rations. Overall,
increasing corn silage intake increased milk concentration of vitamin B12
(Chassaing et al., 2011). Concentrations of vitamin B12 were greater in

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milk of cows receiving a daily supplement of 25 mg of cobalt compared
with an unsupplemented diet, but further increase of daily cobalt supply
from 25 to 75 mg had no effect on milk concentrations of the vitamin
(Akins et al., 2013). A study conducted on 15 commercial dairy farms in
Québec, Canada, showed that vitamin B12 content in milk of cows during
their first two months of lactation varied greatly among farms: from 2.0 to
3.7 ng/g, in spite of small differences in supplemental cobalt among herds
(Figure 1; Duplessis et al., 2011). Recordings of calculated composition of
the rations and analytical measurements were collected, but they did not
allow identifying dietary factors associated with changes in milk concen-
trations of the vitamin. The number of farms involved was possibly too
limited for such survey.
Recently, Rutten et al. (2013) demonstrated that milk concentrations
of vitamin B12 are affected by the genotype of the cow. Genomic regions
associated with milk concentration of the vitamin have been identified,
and this offers an interesting potential for marker-assisted genetic selec-
tion. Nevertheless, data from these few studies highlight that knowledge
on factors affecting milk concentrations of vitamin B12 is very limited. It is
known that vitamin B12 content in milk may be increased by 50% follow-
ing weekly injections of vitamin B12 to dairy cows (Preynat et al., 2009);
in such case, a glass of milk (250 mL) would provide up to 75% of the
RDA. However, in commercial conditions with the variable concentra-
tions of vitamin B12 in milk mentioned above (Duplessis et al., 2011), a
glass of milk from those farms would provide from 20% to almost 40%
(with an average of 33%) of the RDA. Such variations could impact the
relative importance of milk and dairy products as an excellent source of
the vitamin in human nutrition. Therefore, it is important to identify the
factors affecting milk concentrations of vitamin B12 and to develop sus-
tainable nutritional strategies to promote microbial synthesis of this vita-
min in the rumen of the cow and its transfer to milk. Based on the new
findings of Rutten et al. (2013) described previously, genetic selection can
also be a way to increase vitamin B12 in cow milk.
Effect of processing
Raw milk is industrially processed into a wide range of dairy products.
Heating, mechanical or membrane separation, and fermentation are among
the treatments that could potentially alter vitamin B12. Concentrations of

Figure 1. Concentrations of vitamin B12 in milk of cows from 15 dairy herds in the province of Québec, Canada. Adapted from Duplessis et al. (2011).

Apr. 2014, Vol. 4, No. 2 33


Thinkstock/Tom England
vitamin B12 in dairy products range from a low of 1.4 ng/g in butter to
more than 30 ng/g in some cheese varieties (USDA, 2011).
Vitamin B12 was shown to resist pasteurization (75°C for 16 seconds),
and it remains stable during storage of pasteurized milk in a domestic re-
frigerator for nine days (Andersson and Öste, 1994). It is also resistant to
the intense heating treatment (95°C for 5 minutes) applied to milk before
fermentation for yogurt production (Arkbåge et al., 2003). Appreciable
loss (30 to 40%) of vitamin B12 was, however, observed in milk after boil-
ing for 30 minutes or microwave heating for 5 minutes (Watanabe et al.,
1998). Compared with pasteurized milk, vitamin B12 content in canned
evaporated milk is reduced by as much as 65%, despite the concentration
factor due to evaporation (USDA, 2011). Adenosylcobalamin, a predomi-
nant form of vitamin B12 in milk, is known to be sensitive to light, and
Watanabe et al. (2000, 2013) suggested that cobalamin concentration in
milk could be reduced by light exposure. However, no decrease in vita-
min B12 concentration could be detected in milk after exposure to daylight
(Scott et al., 1984) or fluorescent light (Saffert et al., 2006).
Fermentation of heat-treated milk to produce yogurt results in a 25% loss
of vitamin B12 content (Arkbåge et al., 2003). A further decrease (26%) in
vitamin B12 concentration was observed during storage of yogurt at 4°C for
14 days. It has been suggested that the starter culture used for milk fermenta-
tion consumed vitamin B12 and was responsible for these substantial losses
34 Animal Frontiers
(Arkbåge et al., 2003). However, a similar decrease in vitamin B12 concentra-
tion was observed during the storage of non-fermented milk acidified with a
mixture of lactic, acetic, and citric acids (Reddy et al., 1976). This result sug-
gests that the stability of vitamin B12 in acidified milk might be impaired by
low pH or chelating properties of organic acids. Vitamin B12 concentration in
cheese varies from 2.8 ng/g in cream cheese to 8.5 ng/g in cheddar, 22.9 ng/g
in mozzarella, and 33.5 ng/g in Swiss-type cheese (USDA, 2011). Vitamin
B12 is water soluble, and its theoretical concentration in cheese should be less
than that of milk and proportional to moisture content. However, because
vitamin B12 in milk is bound to milk proteins, including caseins (Gizis et al.,
1965), it is partially retained in cheese curd. In hard cheeses, the retention of
vitamin B12 is around 50% (Arkbåge et al., 2003). As for yogurt, cheese stor-
age is likely to influence vitamin B12 concentration due to starter and adjunct
cultures consuming vitamin B12 during ageing. This factor could explain why
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vitamin B12 content is 2.7 times greater in mozzarella than in cheddar cheese.
Mozzarella cheese is consumed a few weeks after production, and the bacte-
rial activity is reduced by cooking and stretching the curd in hot water, while
bacterial activity in cheddar cheese is maintained during a long ripening pe-
riod (up to a few years). Despite a longer ripening period, Swiss-type cheese
contains 45% more vitamin B12 than mozzarella cheese, which is attributed
to the production of vitamin B12 by propionic bacterium during ripening
(Gardner and Champagne, 2005). Propionibacteria are used as an adjunct
culture for manufacture of Swiss-type cheeses where they are responsible
for the characteristic flavor and eye formation (Poonam et al., 2012). In sum-
mary, dairy products are good sources of vitamin B12 and could be even bet-
ter sources by using milk with naturally increased vitamin concentration and
adapting processing conditions to maximize vitamin retention.
The advantage of natural source
In addition to the absolute amount of vitamin B12 present in milk, its
availability for intestinal absorption is also an important factor for assess-
ing the quality of this source of vitamin B12 for humans. Cyanocobala-
min is the synthetic form of vitamin B12 present in most supplements, the
cyanide group being used to stabilize the cobalamin molecule. However,
cyanocobalamin is not biologically active until the cyanide group is enzy-
matically removed (Herbert, 1988). Bioavailability of the synthetic form
of vitamin B12 is inversely dependent on the amount given, being less than
4% in humans and animals receiving prophylactic or therapeutic levels
of supplements (Le Grusse and Watier, 1993; Scott, 1997; Matte et al.,
2010). In milk, vitamin B12 is present as adenosyl-, hydroxo-, and meth-
ylcobalamin (Farquharson and Adams, 1976; Fie et al., 1994). Adenosyl-
cobalamin and methylcobalamin have a coenzymatic activity in mammal
cells and are biologically active whereas hydroxocobalamin is the product
of their photolysis. In a recent experiment (Matte et al., 2012), it was
hypothesized that the important daily provision of vitamin B12 brought
by unprocessed (raw) or processed milk (pasteurized or micro-filtered) is
more efficiently absorbed than the synthetic form (cyanocobalamin) used
in vitamin supplements. Using pigs as an animal model for humans, this
study compared the net portal flux of vitamin B12 (an indicator of intestinal
absorption) after ingestion of milk (raw, pasteurized, or microfiltrated) to
the equivalent amount of cyanocobalamin or to a control diet devoid of
vitamin B12. The efficiency of intestinal absorption of vitamin B12 in milk
is close to 10% regardless of the technological process (raw, pasteurized,
or microfiltered) while the net flux of this vitamin to the portal vein was
undetectable after ingestion of a synthetic supplement of vitamin B12 or a
control meal without vitamin B12 (Table 1).
 Two hypotheses were suggested by Matte et al. (2012) to explain the
greater bioavailability of vitamin B12 naturally present in milk. One was
related to the molecular form of this vitamin in milk, mostly adenosylco-
balamin (Farquharson and Adams, 1976; Fie et al., 1994) as mentioned
previously. Information on the relative intestinal availability of the dif-
ferent forms of cobalamin is scarce. The only data available compared
whole-body retention of crystalline radioactive forms of different cobala-
mins in human subjects (Weissberg and Glass, 1966; Adams et al., 1971).
At doses between 100 and 1000 mg, there was no difference between
cyanocobalamin and hydroxocobalamin (Weissberg and Glass, 1966). At
25 mg, whole-body retention of vitamin B12 was greater after ingestion of
crystalline forms of adenosyl-, hydroxo-, and methylcobalamin than cya-
nocobalamin (Adams et al., 1971). Another explanation for the increased
bioavailability of milk vitamin B12 was related to the presence of specific

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components in milk facilitating its absorption. In pigs, Matte et al. (2010)
reported measurable intestinal absorption of vitamin B12 after ingestion
of cyanocobalamin supplements given in a semi-purified diet, containing
16% of vitamin-free casein (derived from cow milk) whereas Matte et al.
(2012) did not detect any absorption when cyanocobalamin supplements
were mixed with cereals. This finding supports the hypothesis that a milk
component, not destroyed by the technological process for production of
vitamin-free casein, improves intestinal absorption of vitamin B12. In fact,
casein itself could be involved because fractions of this protein were iden-
tified as major components of the protein binding capacity of vitamin B12
in bovine milk (Gizis et al., 1965). According to preliminary data from a
study evaluating methods to increase intestinal absorption of a bolus of
cyanocobalamin infused in the abomasum of dairy cows, absorption of
vitamin B12 in the small intestine was greater when cyanocobalamin was

Thinkstock/moodboard
given in combination with casein than when given alone (Artegoitia et al.,
2013). Food proteins are known to bind vitamin B12 and affect its stability
and bioavailability (Herbert, 1988; Neale, 1990). The binding proteins in
milk are likely to influence acid tolerance and release of vitamin B12 at
gastric pH, providing an adequate protection and release during gastric
transit. Globally, these explanations are in accordance with the numeri-
cally greater, although not statistically significant, efficiency of absorp- ability. In fact, it appears that milk and dairy products could be efficient
tion of dietary 58Co-labeled cyanocobalamin when given in milk (65%) carriers for both endo- and exogenous vitamin B12. Further information
rather than in water or bread (55%; Russell et al., 2001). Moreover, forti- on the effect of milk-specific components on absorption of vitamin B12 is
fied breakfast cereal showed a stronger impact on vitamin B12 status than needed to understand if and how the different milk products (e.g., cheese
meat (Tucker et al., 2000). Since breakfast cereal is usually consumed and yogurt) derived from different fractions of milk retain the original
with milk, proteins from milk might be responsible for increased bioavail- properties of milk in terms of bioavailability of vitamin B12.

Table 1. Intestinal absorption of vitamin B12 in the portal vein Conclusion

of pigs over 24 hours according to sources of vitamin B12.*
The presence of vitamin B12 in cow milk has a considerable impact
Control for the overall worldwide provision of this vitamin for humankind. This
devoid of Cyano- Raw Pasteurized Microfiltered
is all the more important that all plant food sources are devoid of this
Treatments vitamin B12 cobalamin milk milk milk
vitamin. This provision of vitamin B12 from milk is important not only in
Vitamin B12 0 57.5 59.4 71.8 67.8
terms of quantity, as an animal product from ruminants, but also in terms
ingested, µg
of quality, milk vitamin B12 being more bioavailable than its synthetic
Cumulative net ND 0† †
ND 0 5.5 5.6 6.8
form currently available in nutraceutical or pharmaceutical markets. Di-
flux of vitamin B12
to the portal vein etary surveys in human populations showing that vitamin B12 status is
over 24 hours, µg highly correlated with dairy product intake indicate that consumption of
Absorption 0 0 9.2 7.8 10.0 cow milk could become a natural prophylactic tool and then a unique al-
efficiency, % ternative to mandatory fortification of some foods to prevent vitamin B12
* Adapted from Matte et al. (2012). deficiencies in human nutrition.
† ND 0 means that the net flux was not different from 0.

Apr. 2014, Vol. 4, No. 2 35

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About the Authors
J. Jacques Matte is a research scientist
for Agriculture and Agri-Food Canada at
Sherbrooke in Québec. He obtained his
Ph.D. at Université Laval, QC, Canada in
1984, was a post-doc in the UK in 1985,
and had a sabbatical year in France in
1994. He is currently adjunct professor
at Université Laval and at University of
Alberta, AB, Canada. For the last three
years, he was Associate Editor in the non-
ruminant section of Journal of Animal Sci-
ence. His main research areas are related to
metabolism of vitamins and trace minerals
and their impact for growth and for male and female reproduction in pigs.
Correspondence: jacques.matte@agr.gc.ca
Michel Britten has been a senior research
scientist at the Food Research and Devel-
opment Centre, Agriculture and Agri-Food
Canada, at St-Hyacinthe, Québec, Canada
since 1987. He is also an adjunct profes-
sor at Laval University (Food Science and
Nutrition Department) and a member of
the Dairy Science and Technology Cen-
tre (STELA) and the Institute of Nutrition
and Functional Foods (INAF). His main
research area relates to the physical chem-
istry of milk and dairy products. Recent
activities covered the following topics: 1)
development of nutritionally enhanced dairy products, 2) nutrient protection
during gastrointestinal transit, and 3) production of value-added ingredients
derived from cheese whey.
Christiane L. Girard has been a research
scientist at the Dairy and Swine Research
and Development Centre, Agriculture and
Agri-Food Canada, at Sherbrooke, Qué-
bec, Canada since 1985. Her two major
research interests are 1) defining B vitamin
requirements of high-producing dairy cows
to optimize their well-being and metabolic
efficiency, with a current focus on the meta-
bolic interactions between folic acid and vi-
tamin B12 and 2) defining dietary conditions
affecting ruminal synthesis and supply of
B vitamins to dairy cows. For the last three
years, she was the Editor-in-Chief of Canadian Journal of Animal Science and
a member of the board of the Canadian Society of Animal Science.
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