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22.2

SUMMARY 22.1 Overview of Nitrogen Metabolism

The molecular nitrogen that makes up 80% of

the earths atmosphere is unavailable to most
living organisms until it is reduced. This fixation
of atmospheric N2 takes place in certain freeliving bacteria and in symbiotic bacteria in the
root nodules of leguminous plants.

The nitrogen cycle entails formation of

ammonia by bacterial fixation of N2,
nitrification of ammonia to nitrate by soil
organisms, conversion of nitrate to ammonia by
higher plants, synthesis of amino acids from
ammonia by all organisms, and conversion of
nitrate to N2 by denitrifying soil bacteria.

Biosynthesis of Amino Acids

Organisms vary greatly in their ability to synthesize

the 20 common amino acids. Whereas most bacteria and
plants can synthesize all 20, mammals can synthesize
only about half of themgenerally those with simple
pathways. These are the nonessential amino acids,
not needed in the diet (see Table 181). The remainder, the essential amino acids, must be obtained from
food. Unless otherwise indicated, the pathways for the
20 common amino acids presented below are those operative in bacteria.
Glucose

Glucose 6-phosphate

Fixation of N2 as NH3 is carried out by the

nitrogenase complex, in a reaction that
requires ATP. The nitrogenase complex is
highly labile in the presence of O2.
In living systems, reduced nitrogen is
incorporated first into amino acids and then
into a variety of other biomolecules, including
nucleotides. The key entry point is the amino
acid glutamate. Glutamate and glutamine are
the nitrogen donors in a wide variety of
biosynthetic reactions. Glutamine synthetase,
which catalyzes the formation of glutamine
from glutamate, is a main regulatory enzyme of
nitrogen metabolism.
The amino acid and nucleotide biosynthetic
pathways make repeated use of the biological
cofactors pyridoxal phosphate, tetrahydrofolate,
and S-adenosylmethionine. Pyridoxal phosphate is required for transamination reactions
involving glutamate and for other amino acid
transformations. One-carbon transfers require
S-adenosylmethionine and tetrahydrofolate.
Glutamine amidotransferases catalyze reactions
that incorporate nitrogen derived from
glutamine.

22.2 Biosynthesis of Amino Acids

All amino acids are derived from intermediates in glycolysis, the citric acid cycle, or the pentose phosphate
pathway (Fig. 229). Nitrogen enters these pathways by
way of glutamate and glutamine. Some pathways are
simple, others are not. Ten of the amino acids are just
one or several steps removed from the common metabolite from which they are derived. The biosynthetic pathways for others, such as the aromatic amino acids, are
more complex.

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4 steps
Ribose 5phosphate

4 steps

Histidine

Erythrose 4phosphate

Tryptophan
Phenylalanine
Tyrosine

3-Phosphoglycerate

Serine

Phosphoenolpyruvate

Glycine
Cysteine

Pyruvate

Alanine
Valine
Leucine
Isoleucine

Citrate

Oxaloacetate

-Ketoglutarate

Aspartate

Glutamate

Asparagine
Methionine
Threonine
Lysine

Glutamine
Proline
Arginine

FIGURE 229 Overview of amino acid biosynthesis. The carbon

skeleton precursors derive from three sources: glycolysis (pink), the
citric acid cycle (blue), and the pentose phosphate pathway (purple).