BLBS102-c24 BLBS102-Simpson March 21, 2012 13:47 Trim: 276mm X 219mm Printer Name: Yet to Come
24 Chemistry and Biochemistry of Milk Constituents 447
Lactose in Fermented Dairy Products
The fermentation of lactose to lactic acid, by lactic acid bacte-
ria (LAB) is a critical step in the manufacture of all fermented
dairy products (cheese, fermented milks and lactic butter). The
fermentation pathways are well established (see Cogan and Hill
1993, Poolman 2002). Lactose is not a limiting factor in the man-
ufacture of fermented dairy products; only approximately 20%
of the lactose is fermented in the production of these products.
Individuals suffering from lactose intolerance may be able to
consume fermented milks without ill-effects, possibly because
LAB produceβ-galactosidase and emptying of the stomach is
slower than that for fresh milk products, thus, releasing lactose
more slowly into the intestine.
In the manufacture of cheese, most (96–98%) of the lactose
is removed in the whey. The concentration of lactose in fresh
curd depends on its concentration in the milk and on the mois-
ture content of the curd and varies from approximately 1.7%,
w/w, in fresh Cheddar curd to approximately 2.4%, w/w, in fresh
Camembert. The metabolism of residual lactose in the curd to
lactic acid has a major effect on the quality of mature cheese
(Fox et al. 1990, 2000). The resultant lactic acid may remain es-
sentially unchanged in the cheese during ripening (e.g., Cheddar
cheese) or may be catabolised to other compounds, for example
CO 2 and H 2 O, by surface mould in Camembert, or to propionic
acid, acetic acid, H 2 O and CO 2 in Emmental-type cheeses. Ex-
cessive lactic acid in cheese curd may lead to a low pH and a
number of defects, such as a strong, acid, harsh taste, an increase
in brittleness and a decrease in firmness. The pH of full-fat Ched-
dar is inversely related to the lactose/lactic acid content of the
curd. Excess residual lactose may also be fermented by hetero-
fermentative lactobacilli, with the production of CO 2 leading to
an open texture.
In the manufacture of some cheese varieties, for example
Dutch cheese, the curds are washed to reduce the lactose content
and thereby regulate the pH of the pressed curd at approximately
5.3. For Emmental, the curd-whey mixture is diluted with water
by approximately 20%, again to reduce the lactose content of
the curd, maintain the pH at approximately 5.3, and keep the
calcium concentration high, which is important for the textural
properties of this cheese. For Cheddar, the level of lactose, and
hence lactic acid, in the curd is not controlled. Hence, changes
in the concentration of lactose in milk, such as those occurring
throughout lactation, can result in marked changes in the quality
of such cheeses. To overcome seasonal variations in the lactose
content of milk, the level of wash water used for Dutch-type
cheeses is related to the concentrations of lactose and casein
in the milk. Ideally, the lactose-to-protein ratio in any particular
variety should be standardised, for example by washing the curd,
to minimise variations in the concentration of lactic acid, pH and
the quality of cheese.
Oligosaccharides
Lactose is the principal sugar in milk but the milk of most, if not
all, species also contains oligosaccharides, up to hexasaccha-
rides, derived from lactose (the reducing end of the oligosaccha-
rides is lactose and many contain fucose andN-acetylneuraminic
acid). About 130 oligosaccharides have been identified in hu-
man milk; the milk of elephant, bears and marsupials also con-
tains high levels of oligosaccharides. The oligosaccharides are
considered to be important sources of certain monosaccharides,
especially fucose andN-acetylglucosamine, for neonatal devel-
opment, especially of the brain (Urashima et al. 2001, 2009,
2011).
MILK LIPIDS
Definition and Variability
The lipid fraction of milk is defined as those compounds that are
soluble in non-polar solvents (ethyl/petroleum ether or chloro-
form/methanol) and is comprised mainly of triglycerides (98%),
with approximately 1% phospholipids and small amounts of
diglycerides, monoglycerides, cholesterol, cholesterol esters and
traces of fat-soluble vitamins and other lipids. The lipids occur
as globules, 0.1–20μm in diameter, each surrounded by a mem-
brane, the milk fat globule membrane (MFGM), which serves
as an emulsifier. The concentration of total and individual lipids
varies with breed, individual animal, stage of lactation, mastitic
infection, plane of nutrition, interval between milking and point
during milking when the sample is taken. Among the principal
dairy breeds, Friesian/Holsteins produce milk with the lowest fat
content (∼3.5%) and Jersey/Guernsey the highest (∼6%). The
fat content varies considerably throughout lactation; when syn-
chronised calving is practised, the fat content of bulk Friesian
milk varies from approximately 3% in early lactation to>4.5%
in late lactation. Such large variations in lipid content obviously
affect the economics of milk production and the composition of
milk products, but can be modified readily by natural cream-
ing, centrifugal separation or addition of cream, and hence need
not affect product quality. Milk lipids also exhibit variability
in fatty acid composition and in the size and stability of the
globules. These variations, especially fatty acid profile, are es-
sentially impossible to standardise and hence are responsible
for considerable variations in the rheological properties, colour,
chemical stability and nutritional properties of fat-containing
dairy products.
Fatty Acid Profile
Ruminant milk fat contains a wider range of fatty acids than any
other lipid system – up to 400 fatty acids have been reported
in bovine milk fat; the principal fatty acids are the homologous
series of saturated fatty acids with an even number of C-atoms,
C4:0–C18:0,andC18:1. The outstanding features of the fatty acid
profile of bovine milk fat are a high concentration of short- and
medium -chain acids (ruminant milk fats are the only natural
lipids that contain butanoic acid, C4:0) and a low concentration
of polyunsaturated fatty acids.
In ruminants, the fatty acids for the synthesis of milk lipids
are obtained from triglycerides in chylomicrons in the blood
or synthesised de novo in the mammary gland from acetate or
β-hydroxybutyrate produced in the rumen. The triglycerides in