0013 Proteolysis is a major series of events during the
ripening of Swiss cheese. Proteolysis in Swiss cheese is
catalyzed by enzymes indigenous to the milk and
from the cheese microflora and, to a much lesser
extent, from the coagulant. The principal indigenous
proteinase in milk, plasmin, is particularly important
for proteolysis in Swiss-type cheese. Plasmin (fibrino-
lysin, EC 3.4.21.7) is a serine proteinase with an
alkaline pH optimum that is formed from an inactive
precursor, plasminogen, by limited proteolysis. The
activation of plasminogen and plasmin activity in
milk are controlled by a system of plasminogen acti-
vators, plasmin inhibitors, and inhibitors of plasmi-
nogen activators. Plasmin is a particularly heat-stable
proteinase, and hence its activity relative to other
proteolytic enzymes (especially chymosin) is higher
in Swiss cheese than in other varieties, since it survives
the cooking step. Plasmin acts during ripening to
degrade b-casein to g-caseins; the latter accumulate
to greater levels in Swiss cheese than in most other
varieties. Chymosin, or other enzymes from the co-
agulant, are the most important enzymes for the ini-
tial degradation of the caseins during the ripening of
most internal, bacterially ripened varieties. However,
since chymosin is denatured extensively during the
cooking step of Swiss cheese, its action is less import-
ant in this variety than in varieties such as Cheddar.
0014 Intermediate-sized peptides produced by the action
of plasmin or chymosin are hydrolyzed by proteinases
and peptidases from the cheese microflora. Lactic
acid bacteria are auxotrophic for a number of amino
acids and therefore possess a range of proteolytic
enzymes capable of liberating amino acids from
the caseins. Although less well studied than the lacto-
cocci, thermophilic lactobacilli and streptococci used
as starters in Swiss cheese possess generally similar
proteinase and peptidase systems, including a cell
envelope-associated proteinase that is responsible
for degrading many of the intermediate-sized peptides
produced by plasmin and chymosin to shorter pep-
tides. Lactic acid bacteria, including those used as
starters in Swiss cheese, possess a wide range of intra-
cellular peptidases. These enzymes are released into
the cheese matrix following cell lysis, which occurs
during ripening, and are essential for the liberation of
amino acids (AA). Pr. freudenreichii is weakly proteo-
lytic, although it does possess intracellular peptidases
that contribute to ripening.
0015 Proteolysis results ultimately in the liberation of
AA. AA contribute directly to the background flavor
of cheese; some are sweet, some are sour, and many
are bitter. However, the major role of AA in the
development of cheese flavor is now thought to be
as precursor molecules for a range of poorly under-
stood catabolic reactions that lead to the formation of
many volatile flavor compounds. Although amino
acid catabolism has not been studied extensively in
Swiss cheese, it is highly likely that it follows the same
general pathways as in other internal, bacterially
ripened varieties such as Cheddar and Gouda.
Amino acid catabolic reactions include deamination
(to yield NH
3
and a-keto acids), decarboxylation (to
yield amines), formation of alkyl pyrazines, and ca-
tabolism of sulfur-containing and aromatic amino
acids.
Other Swiss-type Cheeses
0016Although Emmental is the Swiss-type variety pro-
duced in the largest amount world-wide, there are
several related Swiss cheese varieties. Gruye
`
re is a
popular Swiss-type cheese that differs from Emmental
in being smaller, with a somewhat stronger flavor and
fewer eyes, and is characterized by the development
of a surface flora. This surface flora (similar to that
which develops on smear-ripened varieties such as
Tilsit and Limburger) is encouraged by ripening for
2–3 weeks at 10
C and for 2–3 months at 15–20
C
and 90–95% ERH. During the hot-room stage, the
cheeses are rubbed with a brine-soaked cloth. Further
ripening at 12–15
C is required before retail; the
cheese is ripe in 8–12 months. Varieties similar to
Gruye
`
re include Raclette (which is manufactured
from raw milk and is acidified by the natural flora
of the milk) and Gruye
`
re de Comte
´
, which is pro-
duced in eastern France. Comte
´
has smaller eyes
than Emmental as a consequence of a low ripening
temperature (c.18
C). The surface of this cheese is
covered by an orange smear (‘morge’); cell numbers at
the surface are typically 10
10
cfu cm
2
, and the micro-
flora is very complex, consisting of coryneform
bacteria, micrococci, and yeasts. Beaufort is a French
variety that is similar to, but larger than (c. 45 kg),
Gruye
`
re and usually develops fewer eyes (as a result
of the growth of mesophilic starter acting on citrate at
a low ripening temperature) than Gruye
`
re. Appenzel-
ler, which originated in Switzerland, is a small cheese
(c. 30 cm) with a soft texture and propionic acid
fermentation that results in the development of a
few eyes. The curds are cooked at 43–45
C. Appen-
zeller is immersed in cider or spiced wine or coated
with a mixture of salt and spices during ripening,
which imparts a distinctive flavor to the cheese.
Maasdammer, a variety developed recently in The
Netherlands, is characterized by the use of a meso-
philic starter and extensive propionic acid fermenta-
tion, which gives large eyes and a domed appearance
to the cheese wheels. Jarlsberg is a Swiss-type cheese
produced in Scandinavia, which is characterized by
large eyes.
1092 CHEESES/Cheeses with ‘Eyes’