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Nutrition in mammary gland health and lactation: Advances over eight Biology of

Lactation in Farm Animals meetings

A. Baldi, F. Cheli, L. Pinotti and C. Pecorini
J ANIM SCI 2008, 86:3-9.
doi: 10.2527/jas.2007-0286 originally published online July 20, 2007

The online version of this article, along with updated information and services, is located on
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Nutrition in mammary gland health and lactation: Advances over eight

Biology of Lactation in Farm Animals meetings1
A. Baldi,2 F. Cheli, L. Pinotti, and C. Pecorini
Department of Veterinary Science and Technology for Food Safety, University of Milan, Italy

neonates and adults. In the past, attempts to probe

the relationship among nutrition, animal health, and
animal products were limited to administering single
dietary components and investigating their biochemical
and metabolic effects. Today, we have genomics, proteomics, and related technologies that allow us to pursue
more holistic investigational strategies. These new
technologies are providing new insights into interactions among nutrition, lactation, and product quality.
The aim of this paper is to review advances in nutritional support of mammary gland function and health
as presented in 14 yr of Biology of Lactation in Farm
Animals (BOLFA) meetings.

ABSTRACT: Over the years, numerous studies have

investigated the mechanisms controlling nutrient
availability and metabolism in the mammary gland and
how dietary interventions can influence these processes. The development of in vivo and in vitro systems
made it possible to explore the trafficking and metabolic
fate of nutrients and how these are influenced by hormones. To improve the quality and safety of milk products, attention has focused on improving animal health
in general and mammary gland health in particular and
also on enhancing the milk content of natural bioactive
milk components that promote the health of human

Key words: nutrition, mammary gland, farm animal, lactation

2008 American Society of Animal Science. All rights reserved.


J. Anim. Sci. 2008. 86(Suppl. 1):39


between nutrition and development, function, and

health of the mammary gland.
The topics explored in the 8 BOLFA meetings include
precursors of and metabolic support for lactation; uptake and metabolism of nutrients by the mammary
gland; manipulation of milk components; and interactions among genes, nutrition, and the endocrine system.
The role of milk as means of delivering nutrients and
bioactive compounds in promoting human and neonate
health also was addressed. Thus, these meetings have
provided a series of snapshots documenting the development of research in nutrition and lactation over the
past 2 decades, with particular attention on the comprehension of basic mechanisms underpinning mammary
gland function. This paper reviews aspects of nutrition
and lactation, as examined at the previous BOLFA
meetings, focusing on dairy cows; however, many of the
concepts discussed are applicable to other farm

Over the years, numerous studies have been presented in the Biology of Lactation in Farm Animals
(BOLFA) workshops on the mechanisms controlling
nutrient availability and metabolism in the mammary
gland, and how dietary interventions can influence
these processes. The first BOLFA workshop took place
in Madrid in 1992, and subsequent workshops have
been held every 2 yr since then (Figure 1). The event
serves as a forum for presenting the results of basic
and applied research on the mammary gland so as to
increase understanding of lactation in farm animals
and has been successful in encouraging the free exchange of scientific ideas on a broad range of topics
centering on the biology of the mammary gland. Scientific progress has been rapid since 1992, and BOLFA
meetings have reflected that progress. A key issue
throughout all the meetings has been the relationship



Presented at the Eighth International Workshop on the Biology

of Lactation in Farm Animals held in Pirassununga, Brazil, August
2123, 2006.
Corresponding author: antonella.baldi@unimi.it
Received May 21, 2007.
Accepted July 17, 2007.

Initially, the studies presented on the topic of supply

of metabolic precursors to support lactation described
mainly the uptake, utilization efficiency, and metabolism of amino acids by the mammary gland, and quanti3

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Baldi et al.

Figure 1. Locations and dates of the Biology of Lactation in Farm Animals (BOLFA) workshops.
fied input-output relationships between nutrient availability and protein secretion. One aim of those studies
was to increase milk protein content by using dietary
manipulation to increase the amino acid supply to the
mammary gland (Murphy and OMara, 1993). In fact,
feeding greater amounts of concentrates increases milk
protein concentration and milk yield. Infusion of casein
or essential amino acids postruminally also increases
milk protein concentration, but responses to feeding
rumen-protected lysine and methionine have been variable and dependent on the stage of lactation, as presented by Rulquin (Rulquin et al., 1993) at the second
BOLFA workshop in Minneapolis.
Metcalf and coworkers (1994) found that short-term
udder infusion of amino acids in lactating dairy cows
resulted in greater milk removal (i.e., 55 to 73%) from
the infused mammary gland than from the noninfused
gland. Those results indicated that mammary gland
uptake of amino acids was influenced by blood levels
of amino acids, although their subsequent fates were
not entirely clear. Some information on the fate of
amino acids in the mammary gland was provided by
studies in goats that indicated that net uptake of amino
acids available for protein synthesis (and therefore, not
oxidized) by the mammary gland, were, in decreasing
order, methionine, phenylalanine, threonine, histidine,
and lysine (Bequette et al., 1994). This order of amino
acid availability has been suggested as an indicator of
amino acid priority for milk protein synthesis.

Although studies on the amino acids, which are limiting for milk production, have focused mainly on essential amino acids, the potentially limiting influence of
nonessential amino acids also was investigated (Meijer
and van der Koelen, 1994). Glutamine, in particular,
has been studied both for its contribution to milk proteins, and for its multiple roles in the body. Glutamine
and glutamate are the major amino acids of milk proteins, constituting approximately 20% of casein amino
acids. Two other features of ruminant glutamine metabolism are important in this context: 1) the low capacity of ruminants to synthesize glutamine compared
with monogastric animals and 2) the response of plasma
and tissue glutamine pools to conditions of metabolic
stress, including high milk production, which resemble
those of most essential amino acids (Meijer et al., 1993).
In the postpartum dairy cow, the rapid increase in milk
yield, simultaneous increase in gut tissue growth, and
the need to synthesize glucose may lead to glutamine
deficiency, particularly in the early postcalving period,
and thereby constitute a limitation on milk yield (Meijer
et al., 1995).
At the 1998 BOLFA workshop in Denver, Colorado,
Petitclerc et al. (2000) examined the genetic relationship among milk yield, feed intake, and feed efficiency;
discussed how rumen function (i.e., carbohydrate digestion, vitamin B requirements, and fatty acid biohydrogenation) could be manipulated to influence milk yield
and composition; and discussed the importance of glu-

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Nutrition in mammary gland health

cose, which is essential for milk synthesis in both ruminants and nonruminants. In particular, postruminal
glucose supply, which enhances glucose absorption, can
balance glucose utilization by portally drained viscera,
sparing endogenous glucose and glucose precursors
and, therefore, increasing total glucose availability to
the rest of the body, including the lactating mammary gland.
At the 2000 BOLFA meeting in The Hague, Hanigan
et al. (2001) examined the various models proposed to
predict milk protein yield. Important components of
these models are mammary gland amino acid metabolism that is regulated by amino acid supply and other
factors, such as energy metabolism, regulation of amino
acids uptake, and mammary blood flow. The secretory
side of the model also must be represented and requires
terms for amino acid oxidation, as well as for protein
synthesis. Hanigan et al. (2001) concluded that a multisubstrate Michaelis-Menten equation form is more consistent with experimental observations and appears to
yield better predictions than single-limiting models.
From these studies, it was evident that although the
amino acid supply influences milk protein concentration and yield (kg of protein/d), the transfer efficiency
of dietary protein to milk is low. Probably this low efficiency of dietary protein to milk is a major factor accounting for the inability of diet to markedly alter milk
protein content, as also reviewed recently by Jenkins
and McGuire (2006). However, studies with cows under
hyperinsulinemic-euglycemic clamp showed that mammary amino acid extraction can be adjusted to enhance
milk protein secretion (McGuire et al., 1994). This finding indicates that substrate uptake from the blood can
be responsive to changes in arterial amino acid concentrations, mammary blood flow, and metabolic activity.
Milk fat content is much more susceptible to dietary
manipulation in relation to the origin of milk fatty acids
in dairy ruminants. Approximately 50% of short- and
medium-chain fatty acids (C4:0 to C16:0) in milk arise
from de novo synthesis in the mammary gland from
acetate and -hydroxybutyrate of ruminal origin,
whereas one-half the palmitic acid and most of the longchain fatty acids of milk arise from uptake into the
mammary gland of lipids from blood. In cows, diets rich
in concentrates, vegetable oils, or fish oil, and those
diets characterized by small particle size can induce
major milk fat depression. By contrast, an increase in
milk fat content occurs when encapsulated lipids are
fed (Chilliard et al., 2001). However, these changes in
milk fatty acid content are generally accompanied by
a major modification in fatty acid profile. Chilliard and
coworkers (2001), in the fifth BOLFA workshop, summarized the effects of dietary factors on milk fat secretion and composition, concentrating on the ability of
different diet formulations to decrease milk fat content
or enhance unsaturated fatty acid concentration to obtain healthier milk for human consumption (see subsequent discussion).

Other topics covered at BOLFA meetings were the

roles of metabolic hormones in regulating milk synthesis, including the effect of insulin on milk protein and
fat yields, and the modulation exerted by somatotropin
and insulin-like growth factors (Flint et al., 2001;
Knight, 2001). Insulin has acute effects on adipose lipogenesis (stimulatory) and lipolysis (inhibitory), but the
ruminant mammary gland is unresponsive to changes
in circulating insulin, which has no apparent effect on
glucose uptake or utilization by the mammary gland.
By inhibiting adipose lipolysis, insulin seems to limit
the supply of milk fat precursors available to the udder,
thus supporting the glucogenic-insulin theory as a
cause of milk fat depression (Bauman and Griinari,
2001). With respect to increases in milk proteins, Petitclerc et al. (2000) suggested that insulin can act directly on either mammary epithelial cell proliferation
or amino acids transport systems in the mammary cells
or indirectly via increases in IGF-I that involve ST.
Somatotropin exerts a systemic effect on all types of
nutrients repartitioning them toward the udder, and
also on the local production of IGF in the mammary
gland (Capuco et al., 2003). The effects of ST administration depend on nutritional status, and when nutritional status is excellent, it produces a substantial increase in milk yield. Specific changes induced by ST
via IGF include greater milk synthesis by secretory
cells and enhanced survival of secretory cells. Insulinlike growth factor-I was proposed to be a cell survival
factor necessary to prevent apoptotic death, as well as
being involved in remodeling events of mammary gland
involution (Knight and Wilde, 1993). It may therefore
be possible to use ST to extend lactation, as was discussed in Lillehammer (Knight, 1997; van Amburgh et
al., 1997) and by Capuco et al. (2003) in Quebec City.
The influence of nutrient supply on lactation also
has been investigated in other species, including sows
(Hartmann et al., 1997; Farmer and Srensen, 2001;
Pere`z Laspiur and Trottier, 2001), ewes (Baldi et al.,
1997), goats (Knight, 1997; Wilde et al., 1997), and
mares (DellOrto et al., 1994a,b; Deichsel and Aurich,
2005). Metabolic mechanisms relating nutrient intake
and lactation performance in the sow were discussed
in several BOLFA workshops. In particular, lipid metabolism in adipose tissue and dietary amino acids, particularly lysine, were shown to be key factors influencing milk production in sows (Pettigrew et al., 1993). In
this field, further progress in amino acid nutrition to
maximize genetic potential for litter weight gain and
milk production of sows was provided by Trottier and
Guan (2000). In discussing amino acid requirements of
the sow, those authors indicated that it is essential not
only to identify which amino acids pools are significant
for the mammary system, but also to understand nutrient interactions in order to obtain an optimal response.
Farmer and Srensen (2001) highlighted the importance of optimal nutrition in prepubertal gilts as a factor influencing mammary development and subsequent
milk production. Those authors also found that high

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Baldi et al.

energy intake by sows during gestation may have detrimental effects on mammary development and milk production, whereas dietary protein had limited effects on
mammary development but could increase subsequent
milk production.


Most of the results obtained in the 1990s in the area
of nutrients uptake were derived from in vivo experiments, particularly lactation trials and arteriovenous
difference studies. Although useful information was obtained from arteriovenous difference studies, they provided little insight into how intracellular nutrients metabolism and trafficking are regulated in the mammary
gland and cannot easily address some aspects such as
mammary cell changes during puberty, pregnancy, and
lactation or the influence of hormones on mammary
development, mammary involution, or nutrient utilization for milk synthesis (Wilde et al., 1997; Purup et
al., 2000).
Studies on secretory pathways within mammary cells
led to important advances in our understanding of the
biology of lactation. In the Hague meeting in 2000,
Boisgard et al. (2001) noted that protein milk composition results from two main processes; the expression of
milk protein encoding genes in the mammary epithelial
cells and the transfer of plasma-borne proteins. An anterograde secretory pathway is responsible for the secretion of newly synthesized proteins in milk, such as
caseins and whey proteins. Following this pathway, the
proteins initially appear on the endoplasmic reticulum,
transiently associate with elements of the Golgi complex, and then concentrate in post-Golgi secretory vesicles before being exported into milk. Beside this main
anterograde transport, a transcytotic pathway is responsible for the transfer of bioactive components (e.g.,
hormones, growth factors, and immunoglobulins) from
the blood or from stroma cells into milk (Boisgard et
al., 2001). In vitro studies using mammary explants,
immortalized cell lines, barrier systems, and alveolilike structures were vital for elucidating these aspects
of uptake and trafficking. In vitro experimental models
of milk secretion also were appraised for investigating
nutrient roles beyond that of supporting milk component synthesis. In particular, they were used to monitor
the effects of interventions to alter the quality or composition of nutrient output during lactation (Clegg et al.,
2001). An increasingly wide variety of biotechnological
tools are now becoming available to elucidate the complex physiology of the mammary gland. Microarrays
for transcriptional profiling and proteomic analyses for
identifying protein expression patterns are useful technologies to investigate changes in the set of genes expressed in the mammary gland and in the proteome
during different physiological stages. These approaches
showed that a huge numbers of genes are differentially
expressed in mammary epithelial cells during preg-

nancy, lactation, and involution. However, cellular biological approaches through in vivo and dynamic studies
are necessary to elucidate the precise links between
gene expression and protein synthesis in the mammary
gland (Ollivier-Bousquet and Devinoy, 2005).
Extensive research on lipid uptake, trafficking, and
secretion at the cellular level has been carried out over
the last 20 yr. Much of our knowledge has been obtained
by studying mechanisms of milk fat depression. In the
Hague meeting in 2000, Bauman and Griinari (2001)
presented their biohydrogenation theory of milk fat depression involving the concept that rumen biohyrogenation can produce trans-10, cis-12 CLA, which seems to
be a potent inhibitor of milk fat synthesis. This concept
is supported by data in the cow showing that postruminal infusion of trans-10, cis-12 CLA reduces the abundance in the mammary gland of mRNA for genes involved in fatty acid uptake [i.e., lipoprotein lipase
(LPL), fatty acid transport (i.e., fatty acid binding protein; FABP), de novo fatty acid synthesis (i.e., acetylCoA carboxylase; ACC and fatty acid synthase; FAS),
desaturation (i.e., stearoyl-CoA desaturase; SCD), and
triglyceride synthesis (i.e., acylglycerol phosphate acyl
transferase; AGPAT and glycerol phosphate acyl transferase; GPAT)]. Levels of trans-10, cis-12 CLA in milk
fat were also found to correlate closely with the decrease
in ACC transcript, and less closely, but still significantly, with the reduction in levels of the enzymes FAS,
LPL, and GPAT, as recently reviewed by these same
authors (Griinari and Bauman, 2006).
Another aspect of the regulation of fatty acid profile
of milk by the mammary gland relates to unsaturated
fatty acids. Desaturase activity in the mammary cells,
that introduces a 9 double bond in the cis conformation, not only converts stearic acid arising from ruminal
biohydrogenation to oleic acid that is secreted in milk,
but also is involved in the synthesis of CLA isomers
in the mammary gland (Bauman and Griinari, 2001;
Chilliard et al., 2001). However, the availability of substrates in the diet is the main factor influencing the
content and profile of milk fatty acids, affecting the
expression of various lipogenic genes at the level of
the mammary gland, as also presented in the latest
ruminant physiology symposium in Denmark (Bernard
et al., 2006).


Milk has long been recognized as a source of macroand micronutrients, immunological components, and
biologically active substances, which not only allow
growth but also promote health in mammalian newborns. At the third BOLFA meeting in Lillehammer in
1996, a workshop section was dedicated to bioactive
components of milk (Zinn, 1997). Many milk proteins,
lipids, lipid-soluble substances, and their digested products are bioactive, including peptides, triacylglycerols,
diacylglycerols, saturated and polyunsaturated fatty

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Nutrition in mammary gland health

acids, phospholipids, vitamins, and vitamin-like substances. Milk also contains protein hormones from the
anterior pituitary (i.e., prolactin and ST), hypothalamus (i.e., ST releasing hormone and somatostatin) and
gut (i.e., vasoactive intestinal peptide, gastrin, and substance P), numerous growth factors (including IGF-I
and IGF-II, IGFBP, epidermal growth factor, transforming growth factors), prostaglandins E and F, lactoferrin, and transferrin (Baumrucker and Blum, 1993;
Meisel, 1997; Schanbacher et al., 1997; Weaver, 1997;
Zinn, 1997).
Schanbacher et al. (1997) reviewed the origins of bioactive peptides in milk, their biochemical properties,
potential as nutraceuticals, and potential pharmacological applications. For example, lactoferrin is a multifunctional protein involved in immunoregulation, antiinflammation, and especially iron metabolism. The biochemical and other properties of lactoferrin were
discussed at the 1998 BOLFA meeting in Denver by
Neville et al. (2000). Those authors noted that lactoferrin binding sites have been found in many cell types
including intestinal cells, hepatocytes, mammary epithelial cells, and platelets. At the 2004 meeting in Bled,
Baumrucker (2005) presented results showing that bovine lactoferrin is involved in the entry of IGFBP-3 into
the nucleus of mammary cells.
At the 2000 BOLFA meeting in the Hague, the roles
of IGF-I and IGFBP in milk were considered by Sejrsen
et al. (2001) who noted that colostrum had greater mitogenic activity than did mature milk due to its high IGFI levels; they proposed that the mitogenic activity of
colostrum was important for mammary gland development, as well as for the developing neonate. Emphasis
at this meeting was also placed on nutritional approaches to alter milk composition for the benefit of
human health (Bauman and Griinari, 2001; Chilliard
et al., 2001; Clegg et al., 2001). In particular, ways of
increasing the polyunsaturated fatty acid content of
milk were all considered (Bauman and Griinari, 2001;
Chilliard et al., 2001) to thereby produce functional
milks. At the most recent BOLFA meeting (2006, Pirassununga, Brazil), the issue of the CLA content of milk
was taken up again (da Silva et al., 2006; Paschoal et
al., 2006). As noted above, mammary lipogenic gene
expression (and hence milk fat synthesis) in cows and
goats seems to be regulated by trans fatty acids, with
trans-10 C18:1 and trans-10, cis-12 CLA emerging as
the main mediators of a milk fat-depressing effect. However, the molecular mechanisms involved in the nutritional regulation of gene expression have not been elucidated completely. Thus, although animal feeding regimens can increase the polyunsaturated lipid content of
milk to make it healthier, the milk thereby becomes
more vulnerable to oxidation, and this stimulated much
interest in milk antioxidants and their transfer from
dietary components in dairy cows, as reviewed extensively at the seventh BOLFA workshop in Bled (Baldi,
2005; Debier et al., 2005; Meglia et al., 2005).

In addition to their antioxidant roles that will be

discussed in the next section, vitamin E and other fatsoluble vitamins and provitamins (e.g., beta-carotene)
are important nutrients per se, for which milk is an
important delivery system. Debier et al. (2005) presented results on the roles of vitamins A and E in the
early stages of life. Vitamin E is necessary to protect
the newborn against oxidative stress, whereas vitamin
A is required for growth and development. Both vitamins are essential for immune system development.
These vitamins must therefore be provided to neonates
in adequate amounts. Colostrum contains relatively
high concentrations of vitamins A and E but mature
milk contains much less. The transfer of these vitamins
into milk does not seem to be simply a passive one
associated with lipid transfer. Vitamin supplementation of gestating and lactating animals appears to increase levels of both vitamin E and A in milk and in
neonatal serum. However, positive effects on young animals are difficult to document. Studies on seals have
shed light on important aspects of the transfer of vitamins A and E from mother to offspring, although much
remains to be learned about the metabolism of these
vitamins during lactation (Debier et al., 2005). Administration of the natural vs. synthetic form of vitamin E
can affect bioavailability and may also influence transfer to milk (Meglia et al., 2005). Like other fat-soluble
micronutrients, fat-soluble vitamins are present in the
milk fat fraction, and this has important implications
for bioaccessibility and bioavailability from milk. In
fact, the fat component of milk is a highly effective
delivery system for fat-soluble vitamins.


The importance of nutrition in maintaining health
of the cow during lactation was an important theme
throughout all BOLFA workshops and was specifically
addressed in the 2004 meeting in Bled. Health may
deteriorate during the transition from late pregnancy
to lactation in dairy cows. After parturition, nutritional
requirements increase rapidly as milk production increase; the negative energy balance period extends for
10 to 12 wk (Butler, 2005). A growing body of evidence
indicates that metabolic disorders associated with negative energy balance may impair fertility, immune status, and antioxidant status. During the transition period in dairy cattle, the total antioxidant capacity is
most susceptible to be compromised. Higher oxygen demand for milk synthesis and secretion increases the
production of reactive oxygen metabolites within the
mammary gland. Sordillo (2005) reported that mammary gland cells from periparturient dairy cows produce more cytokines than those from midlactation animals, resulting in transient increases in intracellular
reactive oxygen species and indicating that during lactation onset, mammary gland epithelial cells are exposed to significant levels of free radicals. The resulting

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Baldi et al.

oxidative damage is a possible cause of increased susceptibility to intramammary infection at this time. In
such situations, the availability of antioxidants to the
mammary cells becomes crucial. The antioxidant system is an integrated system, and deficiencies of one
component can affect the antioxidant efficiency of the
others. Nutrition has a major influence on balance of
prooxidants and antioxidants because several antioxidant system components are micronutrients or are dependent on dietary micronutrients. Vitamin E, betacarotene, and selenium are known to be effective dietary antioxidants in ruminants, whereas several others are under investigation and cannot yet be recommended as routine dietary supplements. In this regard,
Baldi (2005) presented results indicating that adequate
vitamin E status is important for maintaining health
and production in dairy cows. Vitamin E also seems to
be crucially involved in immune system function, such
that supplementation with supranutritional levels of
the vitamin E, in some instances, results in improved
immune responses. Vitamin C and other substances
(e.g., glutathione and selenium) that are associated
with antioxidative defenses, may also be important in
maintaining vitamin E status and function and, hence,
vitamin E availability at the cellular level. The extent
of oxidative stress probably also affects tissue consumption of vitamin E.
Although vitamin A does not have all the activities
of classical antioxidants, it seems to play a role in mammary gland immunobiology, remodeling, and polymorphonuclear leukocyte function during the peripartum
period. In vitro studies of the effect of retinoids on function of polymorphonuclear leukocytes indicate that they
directly affect oxidative burst activity and apoptosis,
but not chemotaxis (Meyer et al., 2005). Vitamin A also
has been suggested to play a role in the morphogenesis,
differentiation, and proliferation of the mammary
gland. Retinol and retinoic acid have been reported to
be potent inhibitors of bovine mammary epithelial cell
proliferation in vitro (Cheli et al. 2003). Although the
exact mode of action at the cellular level remains unknown, the main protective effects of retinol and retinoic acids may be due to regulatory effects on the growth
of normal cells by controlling gene expression of several
growth factors.
The metabolic health status effects of folate, vitamin
B12, and vitamin-like choline also was investigated and
it was found that supplies of these micronutrients are
not always sufficient to maximize health and productivity of dairy cows. Supplementation, especially during
early lactation, can improve lactational performance,
metabolic health, and the nutritional quality of milk
(Girard and Matte, 2005; Pinotti et al., 2005). However,
it also has emerged from these studies that our knowledge of interactions between these 3 micronutrients
is incomplete in the dairy cow, and that a nutritional
approach based on the supply and utilization of individual nutrients is inadequate, further indicating the need

to reappraise the requirements for B-complex vitamins

in dairy cows.

In conclusion, in this review, we have identified major
topics about nutrition and lactation in farm animals,
as discussed in the BOLFA workshops. Although some
of these topics are perennial themes of virtually all
BOLFA workshops, it must be emphasized that the
methods and approaches used to investigate them have
become more sophisticated and powerful over the years,
and are expected to develop more rapidly in the future.
Genomics, proteomics and related technologies have
already emerged important tools for investigating molecular responses to nutrients and their implications
for lactation and nutrition in farm animals, exploiting
the basic sequence information provided by genome

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