136 Part II: Water, Enzymology, Biotechnology, and Protein Cross-linking
processing. Readers should visit the IUBMB en-
zyme nomenclature website for the most up-to-date
details on enzyme classification and nomenclature.
CLASSIFICATION AND
NOMENCLATURE OF ENZYMES
GENERALPRINCIPLES
- First principle. Names purporting to be names of
enzymes, especially those ending in -aseshould
be used only for single enzymes, that is, single
catalytic entities. They should not be applied to
systems containing more than one enzyme. - Second principle. Enzymes are classified and
named according to the reaction they catalyze. - Third principle. Enzymes are divided into groups
on the basis of the type of reactions catalyzed,
and this, together with the name(s) of the
substrate(s), provides a basis for determining the
systematic name and EC number for naming
individual enzymes.
COMMON ANDSYSTEMATICNAMES
- The common name (recommended, trivial, or
working name) follows immediately after the EC
number. - While the common name is normally that used in
the literature, the systematic name, which is
formed in accordance with definite rules, is more
precise chemically. It should be possible to
determine the reaction catalyzed from the
systematic name alone.
SCHEME OFCLASSIFICATION ANDNUMBERING
OFENZYMES
The first Enzyme Commission, in its report in 1961,
devised a system for the classification of enzymes
that also serves as a basis for assigning EC numbers
to them. These code numbers (prefixed by EC),
which are now widely in use, contain four elements
separated by periods (e.g., 1.1.1.1), with the follow-
ing meaning:
- The first number shows to which of the six
divisions (classes) the enzyme belongs, - The second figure indicates the subclass,
- The third figure gives the sub-subclass, and
- The fourth figure is the serial number of the
enzyme in its sub-subclass.
- The fourth figure is the serial number of the
The main classes are
- Class 1.Oxidoreductases (dehydrogenases,
reductases, or oxidases), - Class 2.Transferases,
- Class 3.Hydrolases,
- Class 4.Lyases,
- Class 5.Isomerases (racemases, epimerases,
cis-trans-isomerases, isomerases, tautomerases,
mutases, cycloisomerases), and - Class 6.Ligases (synthases).
Class 1. Oxidoreductases
Enzymes catalyzing oxidoreductions belong to this
class. The reactions are of the form AH 2 B A
BH 2 or AH 2 BA BH H. The substrate
oxidized is regarded as the hydrogen or electron
donor. All reactions within a particular sub-subclass
are written in the same direction. The classification
is based on the order “donor:acceptor oxidoreduc-
tase.” The common name often takes the form “sub-
trate dehydrogenase,” wherever this is possible. If
the reaction is known to occur in the opposite direc-
tion, this may be indicated by a common name of
the form “acceptor reductase” (e.g., the common
name of EC 1.1.1.9 is D-xylose reductase). “Oxi-
dase” is used only in cases where O 2 is an acceptor.
Classification is difficult in some cases because of
the lack of specificity towards the acceptor.
Class 2. Transferases
Transferases are enzymes transferring a group (e.g.,
the methyl group or a glycosyl group), from one
compound (generally regarded as donor) to another
compound (generally regarded as acceptor). The
classification is based on the scheme “donor:acceptor
grouptransferase.” The common names are normally
formed as “acceptor grouptransferase.” In many cas-
es, the donor is a cofactor (coenzyme) carrying the
group to be transferred. The aminotransferases con-
stitute a special case (subclass 2.6): the reaction also
involves an oxidoreduction.
Class 3. Hydrolases
These enzymes catalyze the hydrolysis of various
bonds. Some of these enzymes pose problems be-