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DK 3080 CH 1
DK 3080 CH 1
Milk
2006 by Taylor & Francis Group, LLC
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1
Milk: Main
Characteristics
Milk is dened as the secretion of the mammary glands of mammals, its primary
natural function being nutrition of the young. Milk of some animals, especially
cows, buffaloes, goats and sheep, is also used for human consumption, either as
such or in the form of a range of dairy products. In this book, the word milk will
be used for the normal milk of healthy cows, unless stated otherwise. Occasion-
ally, a com-parison will be made with human milk.
This chapter is meant as a general introduction. Nearly all that is mentioned
with the exception of parts of Section 1.2 is discussed in greater detail in other
chapters. However, for readers new to the eld it is useful to have some idea of the
formation, composition, structure, and properties of milk, as well as the variation
including natural variation and changes due to processing that can occur in
these characteristics, before starting on the main text.
1.1 COMPOSITION AND STRUCTURE
1.1.1 PRINCIPAL COMPONENTS
A classication of the principal constituents of milk is given in Table 1.1. The
principal chemical components or groups of chemical components are those present
in the largest quantities. Of course, the quantity (in grams) is not paramount in all
respects. For example, vitamins are important with respect to nutritive value; en-
zymes are catalysts of reactions; and some minor components contribute markedly
to the taste of milk. More information on milk composition is given in Table 1.3.
Lactose or milk sugar is the distinctive carbohydrate of milk. It is a disac-
charide composed of glucose and galactose. Lactose is a reducing sugar.
The fat is largely made up of triglycerides, constituting a very complicated
mixture. The component fatty acids vary widely in chain length (2 to 20 carbon
atoms) and in saturation (0 to 4 double bonds). Other lipids that are present include
phospholipids, cholesterol, free fatty acids, monoglycerides, and diglycerides.
About four fths of the protein consists of casein, actually a mixture of four
proteins:
S1
-,
S2
-, -, and -casein. The caseins are typical for milk and have some
rather specic properties: They are to some extent phosphorylated and have little
or no secondary structure. The remainder consists, for the most part, of the milk
serum proteins, the main one being -lactoglobulin. Moreover, milk contains nume-
rous minor proteins, including a wide range of enzymes.
The mineral substances primarily K, Na, Ca, Mg, Cl, and phosphate
are not equi-valent to the salts. Milk contains numerous other elements in trace
2006 by Taylor & Francis Group, LLC
4 Milk: Main Characteristics
quantities. The salts are only partly ionized. The organic acids occur largely as
ions or as salts; citrate is the principle one. Furthermore, milk has many miscel-
laneous components, often in trace amounts.
The total content of all substances except water is called the content of dry
matter. Furthermore, one distinguishes solids-not-fat and the content of fat in the
dry matter.
The chemical composition of milk largely determines its nutritional value; the
extent to which microorganisms can grow in it; its avor; and the chemical
reactions that can occur in milk. The latter include reactions that cause off-avours.
1.1.2 STRUCTURAL ELEMENTS
Structure can be dened as the geometrical distribution of the (chemical) compo-
nents in a system. It may imply, as it does in milk, that the liquid contains particles.
This can have important consequences for the properties of the system. For
instance, (1) chemical components are present in separate compartments, which
can greatly affect their reactivity; (2) the presence of particles greatly affects some
physical properties, like viscosity and optical appearance; (3) interaction forces
between particles generally determine the physical stability of the system; and (4)
the separation of some components (fat and casein) is relatively easy.
Figure 1.1 shows the main structural elements of milk. Of course, the gure
is schematic and incomplete. Some properties of the structural elements are given
TABLE 1.1
Approximate Composition of Milk
Component
Average Content
in Milk (% w/w)
Range
a
(% w/w)
Average Content in
Dry Matter (% w/w)
Water 87.1 85.388.7
Solids-not-fat 8.9 7.910.0
Fat in dry matter 31 2238
Lactose 4.6 3.85.3 36
Fat 4.0 2.55.5 31
Protein
b
3.3 2.34.4 25
casein 2.6 1.73.5 20
Mineral substances 0.7 0.570.83 5.4
Organic acids 0.17 0.120.21 1.3
Miscellaneous 0.15 1.2
Note: Typical for milks of lowland breeds.
a
These values will rarely be exceeded, e.g., in 1 to 2% of samples of separate
milkings of healthy individual cows, excluding colostrum and milk drawn shortly
before parturition.
b
Nonprotein nitrogen compounds not included.
2006 by Taylor & Francis Group, LLC
1.1 Composition and Structure 5
in Table 1.2, again in a simplied form; the numerical data mentioned are meant
only to dene orders of magnitude. The table clearly shows that aspects of colloid
chemistry are essential for understanding the properties of milk and the many
changes that can occur in it. All particles exhibit Brownian motion; they have an
electrostatic charge, which is negative at the pH of milk. Their total surface area
is large.
Fat globules. To a certain extent, milk is an oil-in-water emulsion. But the
fat globules are more complicated than emulsion droplets. In particular, the
surface layer or membrane of the fat globule is not an adsorption layer of one
FIGURE 1.1 Milk viewed at different magnications, showing the relative size of struc-
tural elements (A) Uniform liquid. However, the liquid is turbid and thus cannot be
homogeneous. (B) Spherical droplets, consisting of fat. These globules oat in a liquid
(plasma), which is still turbid. (C) The plasma contains proteinaceous particles, which are
casein micelles. The remaining liquid (serum) is still opalescent, so it must contain other
particles. The fat globules have a thin outer layer (membrane) of different constitution.
(From H. Mulder and P. Walstra, The Milk Fat Globule, Pudoc, Wageningen, 1974.)
C
50000
B
500
A
5
Milk
Plasma
Fat
globules
Fat
globule
Serum
Membrane
Casein
micelles
2006 by Taylor & Francis Group, LLC
6 Milk: Main Characteristics
single substance but consists of many components; its structure is complicated.
The dry mass of the membrane is about 2.5% of that of the fat. A small part of
the lipids of milk is found outside the fat globules. At temperatures below 35C,
part of the fat in the globules can crystallize. Milk minus fat globules is called
milk plasma, i.e., the liquid in which the fat globules oat.
Casein micelles consist of water, protein, and salts. The protein is casein.
Casein is present as a caseinate, which means that it binds cations, primarily
calcium and magnesium. The other salts in the micelles occur as a calcium
phosphate, varying somewhat in composition and also containing a small amount
of citrate. This is often called colloidal phosphate. The whole may be called
calcium-caseinate/calcium-phosphate complex. The casein micelles are not
micelles in the colloid-chemical sense but just small particles. The micelles have
TABLE 1.2
Properties of the Main Structural Elements of Milk
Milk
Plasma
Serum
Fat Globules Casein Micelles
Globular
Proteins
Lipoprotein
Particles
Main components Fat Casein, water,
salts
Serum protein Lipids, proteins
To be considered as Emulsion Fine dispersion Colloidal
solution
Colloidal
dispersion
Content
(% dry matter)
4 2.8 0.6 0.01
Volume fraction 0.05 0.1 0.006 10
4
Particle diameter
a
0.110 m 20400 nm 36 nm 10 nm
Number per ml 10
10
10
14
10
17
10
14
Surface area
(cm
2
/ml milk)
700 40000 50000 100
Density
(20C; kg m
3
)
920 1100 1300 1100
Visible with Microscope Ultramicroscope Electron
microscope
Separable with Milk separator High-speed
centrifuge
Ultraltration Ultraltration
Diffusion rate
(mm in 1h)
a
0.0 0.10.3 0.6 0.4
Isoelectric pH 3.8 4.6 45 4
Note: Numerical values are approximate averages.
a
For comparison, most molecules in solution are 0.4 to 1 nm diameter, and diffuse, say, 5 mm
in 1 h. 1 mm = 10
3
m = 10
6
nm = 10
7
.
2006 by Taylor & Francis Group, LLC
1.2 Milk Formation 7
an open structure and, accordingly, contain much water, a few grams per gram
of casein. Milk serum, i.e., the liquid in which the micelles are dispersed, is milk
minus fat globules and casein micelles.
Serum proteins are largely present in milk in molecular form or as very small
aggregates.
Lipoprotein particles, sometimes called milk microsomes, vary in quantity
and shape. Presumably, they consist of remnants of mammary secretory cell
membranes. Few denitive data on lipoprotein particles have been published.
Cells, i.e., leukocytes, are always present in milk. They account for about
0.01% of the volume of milk of healthy cows. Of course, the cells contain all
cytoplasmic components such as enzymes. They are rich in catalase.
Table 1.3 gives a survey of the average composition and structure of milk.
1.2 MILK FORMATION
Milk components are for the most part formed in the mammary gland (the udder)
of a cow, from precursors that are the results of digestion.
Digestion. Mammals digest their food by the use of enzymes to obtain simple,
soluble, low-molar-mass components, especially monosaccharides; small pep-
tides and amino acids; and fatty acids and monoglycerides. These are taken up
in the blood, together with other nutrients, such as various salts, glycerol, organic
acids, etc. The substances are transported to all the organs in the body, including
the mammary gland, to provide energy and building blocks (precursors) for
metabolism, including the synthesis of proteins, lipids, etc.
In ruminants like the cow, considerable predigestion occurs by means of
microbial fermentation, which occurs for the most part in the rst stomach or
rumen. The latter may be considered as a large and very complex bio-fermenter.
It contains numerous bacteria that can digest cellulose, thereby breaking down
plant cell walls, providing energy and liberating the cell contents. From cellulose
and other carbohydrates, acetic, propionic, butyric and lactic acid are formed, which
are taken up in the blood. The composition of the organic acid mixture depends
on the composition of the feed. Proteins are broken down into amino acids. The
rumen ora uses these to make proteins but can also synthesize amino acids from
low-molar-mass nitrogenous components. Further on in the digestive tract the
microbes are digested, liberating amino acids. Also, food lipids are hydrolyzed in
the rumen and partly metabolized by the microorganisms. All these precursors can
reach the mammary gland.
Milk Synthesis. The synthesis of milk components occurs for the greater
part in the secretory cells of the mammary gland. Figure 1.2 illustrates such a
cell. At the basal end precursors of milk components are taken up from the blood,
and at the apical end milk components are secreted into the lumen. Proteins are
formed in the endoplasmic reticulum and transported to the Golgi vesicles, in
which most of the soluble milk components are collected. The vesicles grow in
size while being transported through the cell and then open up to release their
contents in the lumen. Triglycerides are synthesized in the cytoplasm, forming
2006 by Taylor & Francis Group, LLC
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TABLE 1.3
Composition and Structure of Milk
a
SERUM
Water 790 g Organic acids Proteins
citrate 1600 mg casein +
Carbohydrates formate 40 mg -lactoglobulin 3.2 g
lactose 46 g acetate 30 mg -lactalbumin 1.2 g
glucose 70 mg lactate 20 mg serum albumin 0.4 g
others oxalate 20 mg immunoglobulins 0.8 g
others 10 mg proteose peptone +
Minerals others
Ca, bound 300 mg Gases Nonprotein nitrogenous
Ca, ions 90 mg oxygen 6 mg compounds
Mg 70 mg nitrogen 16 mg peptides +
K 1500 mg Lipids amino acids 50 mg
Na 450 mg glycerides + urea 250 mg
Cl 1100 mg fatty acids 20 mg ammonia 10 mg
phosphate 1100 mg phospholipids 100 mg others 300 mg
sulfate 100 mg cerebrosides 10 mg Enzymes
bicarbonate 100 mg sterols 15 mg acid phosphatase
others peroxidase
Trace elements many others
Zn 3 mg Vitamins, e.g. Phosphoric esters ~300 mg
Fe 120 g riboavin 2 mg Others
Cu 20 g ascorbic acid 20 mg
many others
a
Approximate average quantities in 1 kg milk. Note: The water in the casein micelles contains some small-molecule solutes.
FAT GLOBULE
triglycerides
diglycerides
monoglycerides
Glycerides
40 g
0.1 g
10 mg
60 mg
60 mg
100 mg
0.3 mg
Fatty acids
Sterols
Carotenoids
Vitamins A, D, E, K
Water
Others
CASEIN MICELLE
casein
proteose peptone
lipase
plasmin
Ca
phosphate
citrate
K, Mg, Na
Protein
Salts
Water
Enzymes
850 mg
1000 mg
150 mg
26 g
+
~80 g
MEMBRANE
LIPOPROTEIN
PARTICLE
lipids
protein
enzymes
water
LEUKOCYTE
Many enzymes
e.g., catalase
Nucleic acids
Water
water +
protein 700 mg
phospholipids 250 mg
cerebrosides 30 mg
glycerides +
fatty acids 15 mg
streols 15 mg
other lipids
enzymes
alkaline phosphatase
xanthine oxidase
many others
Cu 4 g
Fe 100 g
2 g
2006 by Taylor & Francis Group, LLC
1.2 Milk Formation 9
small globules, which grow while they are transported to the apical end of the
cell. They become enrobed by the outer cell membrane (or plasmalemma) while
being pinched off into the lumen. This type of secretion is called merocrine,
which means that the cell remains intact.
Table 1.4 gives some information about the synthesis of specic components.
Most are synthesized in the cell. Others are taken up from the blood but, generally,
not in the same proportion as in the blood; see, especially, the salts. This means
that the cell membranes have mechanisms to reject, or allow passage of, specic
components. Some substances, notably water and small lipophilic molecules, can
FIGURE 1.2 Stylized diagram of a mammary secretory cell. Below is the basal part, on
top the apical part of the cell. The cell is bounded by other secretory cells to form the
glandular epithelium. See text for further details. (From P. Walstra and R. Jenness, Dairy
Chemistry and Physics, Wiley, New York, 1984. With permission.)
Ribosomes
Basement
membrane
Golgi apparatus
Cytosol
Lysosome
Outer cell
membrane
(plasmalemma)
Golgi vesicle
with casein
micelles
Endoplasmic
reticulum
Mitochondrion
Nascent
fat globule
Junctional
complex
Microvillus
5 m
Nucleus
LUMEN
2006 by Taylor & Francis Group, LLC
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TABLE 1.4
Synthesis of Important Milk Components
Milk Component Precursor in Blood Plasma Synthesis of Component
Name
Concentration
(% w/w) Name
Concentration
(% w/w)
In the
Secretory Cell?
Specic
for Milk?
Specic for
the Species?
Water 86 Identical 91 No No No
Lactose 4.7 Glucose
a
0.05 Yes Yes No
Protein
Caseins 2.6
0.32
0.12
0.01
Yes Yes Yes
b
-lactoglobulin Amino acids 0.04 Yes Yes Yes
-lactalbumin Yes Yes Yes
Lactoferrin Yes No Yes
Serum albumin 0.04 Identical 3.2 No No Yes
Immunoglobulins 0.07 Most are identical 1.5 No No Yes
Enzymes Trace Various Yes
c
No
c
Yes
Lipids
Triglycerides 4 Partly Partly
Phospholipids 0.03 Some lipids 0.3
Citric Acid 0.17 Glucose
a
0.05 Yes No No
Minerals Identical No No No
Ca 0.13 0.01
P
d
0.09 0.01
Na 0.04 0.34
K 0.15 0.03
Cl 0.11 0.35
a
Glucose can also be formed in the secretory cells from some amino acids.
b
All proteins are species specic, but comparable proteins occur in the milk of all ruminants.
c
Is not true for all enzymes.
d
In various phosphates.
Acetic acid
-Hydroxy butyric acid
A
0 01
0 006
.
.
ccylglycerols