Can the Replacement of a Single Atom in the Enzyme Horseradish Peroxidase Convert It
into a Monoxygenase? A Density Functional Study
Sam P. de Visser*
Contribution from the Manchester Interdisciplinary Biocenter and the School of Chemical Engineering and
Analytical Science, The UniVersity of Manchester, 131 Princess Street, Manchester M1 7DN, United Kingdom
ReceiVed: August 31, 2006; In Final Form: September 1, 2006
Density functional calculations on horseradish peroxidase mutants are presented, whereby one or two of the
nitrogen atoms of the axial histidine ligand have been replaced by phosphorus atoms. Our calculations show
that phosphorus entices a push effect on the oxoiron group, whereas a histidine side chain withdraws electrons.
As a result, we predict that a phosphorus-substituted histidine ligand will convert the active form of a peroxidase
into a monoxygenase. This subsitution may be useful for the bioengineering of commercially exploitable
enzymes.
Peroxidases are heme enzymes that catalyze the detoxification
of hydrogen peroxide to water and molecular oxygen.
1
The
active species (Compound I, CpdI) of heme peroxidases, such
as horseradish peroxidase (HRP) or cytochrome c peroxidase
(CcP), contains an oxoiron group embedded in a heme.
2
The
peroxidases differ from other heme enzymes such as the
cytochrome P450 enzymes in the nature of the axial ligand
bound to iron, namely the active species in P450 enzymes is
bound to the protein backbone via a thiolate linkage of a
cysteinate residue, whereas peroxidases generally bind to a
histidine side chain.
3
The differences in axial ligand result in
different chemical properties of the active species, i.e., a
cysteinate ligand exerts a push effect on the iron, while histidine
withdraws electron density.
4
As a result, these two ligands give
differences in electronic as well as catalytic properties to the
enzyme active site.
5,6
Attempts to mutate the axial ligand of
CcP into cysteinate were unsuccessful and resulted in oxidation
of the ligand into cysteic acid.
7
In the past we extensively studied
the axial ligand effect on the catalytic properties of oxoiron
porphyrin models using theoretical methods.
4c,6
In line with this,
in this work we present the effect of replacing one or more
nitrogen atoms of the histidine axial ligand by phosphorus atoms
and the changes this exerts on the catalyst.
Recent studies on R-iminophospholide ligands in coordination
chemistry showed extensive charge delocalization from the
phosphorus atom.
8
Moreover, biphosphines were shown to have
strong π-accepting properties and as a result stabilize transition
metals in low oxidation states.
9
To find out what effects the
replacement of a nitrogen atom by a phosphorus atom has on
the electronic and catalytic properties of HRP CpdI, we have
investigated models of CpdI with only the axial bound nitrogen
atom replaced by phosphorus (CpdI(1P)) and a model in which
both nitrogen atoms of the imidazole ring are replaced by
phosphorus (CpdI(2P)).
The model of the active species of peroxidase CpdI is an
oxoiron porphyrin (without side chains) that is bound to an axial
imidazole group, and the overall charge of the model is +1.
All geometries were fully optimized with Jaguar 5.5 using the
UB3LYP hybrid density functional method.
10,11
We employed
an LACVP basis set for the geometry optimizations and ran
single point calculations with an LACV3P+* basis set to
confirm the energetics.
12
All energetics reported in this work
are calculated at the LACV3P+* level of theory with ZPE
corrections at the LACVP level. The charge distributions were
calculated with the Natural Bond Analysis (NBO) program as
implemented in Gaussian-03.
13,14
Figure 1 shows the optimized geometries and group spin
densities (F) of CpdI(1P) and CpdI(2P) in the lowest lying
quartet and doublet spin states. Similarly to CpdI of peroxidase
and P450 models, CpdI(1P) and CpdI(2P) are also described
as a triradicaloid system: two unpaired electrons located in π*
orbitals along the FeO bond and the third one in a nonbonding
heme orbital with a
2u
symmetry.
5,15
These three electrons are
either ferromagnetically coupled into an overall quartet spin state
or antiferromagnetically coupled into a doublet spin state. As
the coupling between the π*
FeO
and a
2u
electrons is weak, the
doublet-quartet energy gap is small: ΔE+ZPE )-0.2 kcal
mol
-1
in CpdI(1P) and +0.1 in CpdI(2P), where a minus sign
implies a doublet spin ground state. These energy differences
are virtually identical to those obtained for HRP CpdI, where
an energy gap of +0.1 kcal mol
-1
was obtained using the same
methods and basis sets. The group spin densities match those
obtained for HRP CpdI closely, the only minor difference being
a slightly higher absolute value on the axial ligand here.
5b
In
P450 models also a significant axial ligand spin density was
obtained due to mixing of the a
2u
orbital with a σ
S
orbital on
the ligand.
4c
It appears that the phosphorus substituted structures
have similar mixing patterns but to a somewhat lesser degree
than thiolate ligated systems.
The optimized geometries of CpdI(1P) and CpdI(2P) are
similar to those obtained for either HRP or P450 CpdI.
5,15
The
Fe-O distances are closer to those obtained for P450 CpdI than
those for HRP CpdI, where values of 1.651 (1.648) Å and 1.621 * Corresponding author. E-mail: sam.devisser@manchester.ac.uk.
20759
2006, 110, 20759-20761
Published on Web 09/27/2006
10.1021/jp065660q CCC: $33.50 © 2006 American Chemical Society