WO2004108946A2 - Crystals of herpes simplex virus 1 dna polymerase and/or herpes simplex virus 1 dna polymerase-like proteins and the use thereof - Google Patents

Abstract

The x-ray crystal structure of Herpes simplex virus 1 DNA polymerase or Herpes simplex virus 1 DNA polymerase-like proteins is useful for solving the structure of other molecules or molecular complexes, and identifying and/or designing potential modifiers of Herpes simplex virus 1 DNA polymerase activity.

Description

CRYSTALS OF HERPES SIMPLEX VIRUS 1 DNA POLYMERASE AND/OR HERPES SIMPLEX VIRUS 1 DNA POLYMERASE-LIKE PROTEINS AND THE USE THEREOF

This application claims the benefit of U.S . Provisional Application No.

60/475,626, filed June 4, 2003, which is herein incorporated by reference in its entirety.

FIELD OF THE INVENTION This invention relates to the crystallization and structure determination of

Herpes simplex virus 1 DNA polymerase, also known as HSN 1 DΝA polymerase or HSV 1 POL.

BACKGROUND Viruses of herpesviridae family cause a wide range of serious diseases in humans. Herpes simplex 1 virus (HSN 1) causes painful facial lesions (e.g., cold sores). HSV 2 infections in humans occur mainly in the genital tract. Epstein- Barr virus (EBV) and Varicella-Zoster virus (VZV) cause infectious mononucleosis in children and shingles (chickenpox) in humans at all ages, respectively. In immuno-compromised patients, cytomegalovirus (CMV) opportunistically infects the gastrointestinal tract and causes life-threatening colitis. Serious eye infections can also cause blindness. Treatments of these infections are important to human health and also have high market value. However, existing antiherpetic drugs (e.g., acyclovir, valaciclovir, famciclovir, ganciclovir, foscarnet and cidofovir) have narrow activity spectra, and drug resistance has been observed with some drugs in immuno-compromised patients. There is a continuing need in the art for safe and effective broad-spectrum antiherpetic drugs.

An important target in treating herpesvirus is the viral DΝA polymerase (POL), since viral replication depends on this protein. These large proteins have multiple enzymatic activities and interact with many other replication proteins, such as viral primase/helicase, to fulfill their functions. It is of great interest to know how Heφes virus POL binds to DNA templates and nucleic acid substrates. The large conformational changes of these proteins during the polymerization reaction are important to understand their catalytic mechanism. One of the most important tools to study protein mechanism is X-ray crystallography. What is needed in the art is a crystal structure of a Herpes virus DNA polymerase.

SUMMARY OF THE INVENTION In one aspect, the present invention provides a crystal of Herpes simplex virus 1 DNA polymerase, and methods of crystallizing a Heφes simplex virus 1 DNA polymerase. Preferably, the crystal has orthorhombic space group symmetry P2₁2₁2₁. Preferably, the crystal includes a unit cell defined by the dimensions a, b, c, α, β, and γ, wherein a is about 80 A to about 130 A, b is about 100 A to about 150 A, c is about 200 A to about 250 A, and =β=γ=90°. Preferably the Heφes simplex virus 1 DNA polymerase is a monomer and the crystal includes two monomers per crystallographic asymmetric unit. Preferably, the Heφes simplex virus 1 DNA polymerase has an amino acid sequence including SEQ ID NO: 1 , and more preferably, consisting of SEQ ID NO: 1. Optionally, the Heφes simplex virus 1 DNA polymerase has an amino acid sequence including SEQ ID NO: 1, with the proviso that a methionine is replaced with selenomethionine. Methods of using such crystals are also provided.

In another aspect, the present invention provides a Heφes simplex virus 1 DNA polymerase having an amino acid sequence including SEQ TDNO:l. Preferably, the Heφes simplex virus 1 DNA polymerase has an amino acid sequence consisting of SEQ TDNO: 1.

In another aspect, the present invention provides a method of preparing a Heφes simplex virus 1 DNA polymerase. Preferably, the Heφes simplex virus 1 DNA polymerase is prepared by expression in insect cells. Preferably a fusion protein is expressed in insect cells. In one embodiment, the fusion protein is a constructed from maltose binding protein and full length Heφes simplex virus 1 DNA polymerase. Preferably, the fusion protein has an amino acid sequence including SEQ J-D NO:2. Preferably, the Heφes simplex virus 1 DNA polymerase is formed by digestion of the fusion protein, and more preferably the fusion protein is digested by an endogenous protease or factor Xa or other specific protease introduced by mutagenesis in the linker region. Preferably, the Heφes simplex virus 1 DNA polymerase has an amino acid sequence including SEQ J-D NO: 1 , and more preferably, consisting of SEQ ID NO: 1.

In another aspect, the present invention provides a method of identifying the ability for a potential modifier to bind to Heφes simplex virus 1 DNA polymerase. In one embodiment, the method includes: exposing a crystal of Heφes simplex virus 1 DNA polymerase to one or more samples including a potential modifier of Heφes simplex virus 1 DNA polymerase; and determining whether a potential modifier-Heφes simplex virus 1 DNA polymerase molecular complex is formed. Preferably, the exposing includes soaking. Optionally, the one or more samples include a plurality of potential modifiers.

In another aspect, the present invention provides a method of acquiring structural information for designing potential modifiers for forming molecular complexes with Heφes simplex virus 1 DNA polymerase. In one embodiment, the method includes: exposing a crystal of Heφes simplex virus 1 DNA polymerase to a library of potential modifiers having diverse shapes; and determining whether a potential modifier-Heφes simplex virus 1 DNA polymerase molecular complex is formed. Preferably, the method further includes identifying the chemical entity that forms the potential modifier upon determination of the formation of the potential modifier-Heφes simplex virus 1 DNA polymerase molecular complex. Preferably, determining and/or identifying includes calculating an electron density function and/or collecting x-ray diffraction data.

In another aspect, the present invention provides a molecule or molecular complex including at least a portion of a Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active site, wherein the active site includes the amino acids listed in Table 2, Table 3, or Table 4, and the active site is defined by a set of points having a root mean square deviation of less than about 3 A from points representing the backbone atoms of said amino acids as represented by the structure coordinates listed in Table 1.

In another aspect, the present invention provides a molecule or molecular complex that is structurally homologous to a Heφes simplex virus 1 DNA polymerase molecule or molecular complex, wherein the Heφes simplex virus 1 DNA polymerase molecule or molecular complex is represented by at least a portion of the structure coordinates listed in Table 1.

In another aspect, the present invention provides a scalable three- dimensional configuration of points, at least a portion of said points derived from structure coordinates as listed in Table 1 of at least a portion of a Heφes simplex virus 1 DNA polymerase molecule or molecular complex that includes a Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase- like active site. Preferably, at least a portion of the points derived from the Heφes simplex virus 1 DNA polymerase structure coordinates are derived from structure coordinates representing the locations of at least the backbone atoms of amino acids defining a Heφes simplex virus 1 DNA polymerase active site including the amino acids listed in Table 2, Table 3, or Table 4. Optionally, the scalable three-dimensional configuration of points is displayed as a holographic image, a stereodiagram, a model, or a computer-displayed image.

In another aspect, the present invention provides a machine-readable data storage medium including a data storage material encoded with machine readable data which, when using a machine programmed with instructions for using said data, displays a graphical three-dimensional representation of a molecule or molecular complex. A useful molecule or molecular complex includes: (i) a molecule or molecular complex including at least a portion of a Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active site including the amino acids listed in Table 2, the active site defined by a set of points having a root mean square deviation of less than about 3 A from points representing the backbone atoms of said amino acids as represented by structure coordinates listed in Table 1 for a Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like molecule or molecular complex; and (ii) a molecule or molecular complex that is structurally homologous to a Heφes simplex virus 1 DNA polymerase molecule or molecular complex that is represented by at least a portion of the structure coordinates listed in Table 1.

In another aspect, the present invention provides a method for obtaining structural information about a molecule or a molecular complex of unknown structure. The method includes: crystallizing the molecule or molecular complex; generating an x-ray diffraction pattern from the crystallized molecule or molecular complex; and applying to the x-ray diffraction pattern at least a portion of the structure coordinates as set forth in Table 1 for Heφes simplex virus 1 DNA polymerase to generate a three-dimensional electron density map of at least a portion of the molecule or molecular complex whose structure is unknown.

In another aspect, the present invention provides a method for homology modeling a Heφes simplex virus 1 DNA polymerase homolog. The method includes: aligning the amino acid sequence of a Heφes simplex virus 1 DNA polymerase homolog with an amino acid sequence of Heφes simplex virus 1 DNA polymerase and incoφorating the sequence of the Heφes simplex virus 1 DNA polymerase homolog into a model of Heφes simplex virus 1 DNA polymerase formed from structure coordinates as set forth in Table 1 for Heφes simplex virus 1 DNA polymerase to yield a preliminary model of the Heφes simplex virus 1 DNA polymerase homolog; subjecting the preliminary model to energy minimization to yield an energy minimized model; and remodeling regions of the energy minimized model where stereochemistry restraints are violated to yield a final model of the Heφes simplex virus 1 DNA polymerase homolog. In another aspect, the present invention provides computer-assisted methods for identifying, designing, or making a potential modifier of Heφes simplex virus 1 DNA polymerase activity. Preferably the methods include screening a library of chemical entities.

ABBREVIATIONS

The following abbreviations may be used throughout this disclosure:

Heφes simplex virus 1 (HSV 1) Heφes simplex virus 1 DNA polymerase (HSV 1 POL)

4-(2-Hydroxyethyl)-l-piperazineethanesulfonic acid (HEPES)

E. coli maltose binding protein (MBP)

2-Amino-2-hydroxymethyl- 1 ,3-propanediol (TRIS)

Polyethylene Glycol (PEG) Dimethyl sulfoxide (DMSO)

Dithiothreitol (DTT)

Multiple anomalous dispersion (MAD)

Root mean square (r.m.s.)

Root mean square deviation (r.m.s. d.) The following amino acid abbreviations are used throughout this disclosure:

A = Ala = Alanine T = Thr = Threonine V = Val = Valine C = Cys = Cysteine L = Leu = Leucine Y = Tyr = Tyrosine I = JJe = Isoleucine N = Asn = Asparagine P = Pro = Proline Q = Gin = Glutamine F = Phe = Phenylalanine D = Asp = Aspartic Acid W = Tφ = Tryptophan

E = Glu = Glutamic Acid M = Met = Methionine K = Lys = Lysine G = Gly = Glycine R = Arg = Arginine S = Ser = Serine H = His = Histidine

BRIEF DESCRIPTION OF THE FIGURES

Figure 1 is an illustration of the amino acid sequence of HSV- 1 strain KOS DNA polymerase (SEQ ID NO: 1 ).

Figure 2 is an illustration of a nucleotide sequence (SEQ ID NO:2) for a gene that encodes HSV-1 strain KOS DNA polymerase.

Figure 3 is an illustration of the amino acid sequence of MBP-HSV-1 strain KOS DNA polymerase fusion protein (SEQ 3D NO:3). Figure 4 is an illustration of a nucleotide sequence (SEQ ID NO:4) for a gene that encodes MBP-HSV-1 strain KOS DNA polymerase fusion protein.

Figure 5 is an illustration of a nucleotide sequence (SEQ ID NO:5) for the pACMAL vector.

Figure 6 illustrates the construction of the pACMAL vector (not to scale). The pMAL-CRI vector (NEBiolabs, Beverly, MA) was digested with Asel. The fragment containing the Mai E sequences and multiple cloning site was gel isolated, and treated with Klenow. The baculovirus expression vector, pAc373 was digested with BamHI and treated with Klenow. The Mai E sequences were then blunt end ligated into the expression vector. Figure 7 is a schematic representation of exemplary restriction nuclease sites of the pACMAL vector.

Figure 8 is a schematic illustration of the construct of the linker between MBP and HSV 1 POL in the fusion protein. Figure 8a is an illustration of an amino acid sequence of the linker with one engineered protease cleavage site (factor Xa site, JEGRI). However, digestion can also be achieved by the endogenous protease(s), producing a product with N43 as the N-terminus (N43- C HSV POL). Since we obtained crystals on N43-C but not with factor Xa digestion product, a second protease digestion site was engineered in front of

N43. Figure 8b illustrates a construct modified from 8a with nucleotides changed from encoding CLRQ to IEGR (factor Xa) (nucleotides 5528-5539 of pACMAL-HSV IPOL reference sequence). Both constructs were used in this application resulting the same crystal form.

Figure 9 is a schematic illustration of an HSV 1 POL purification scheme flow-chart. Straight arrows point to the location of protease digestion to remove

MBP from HSV 1 POL. Figure 10 illustrates a stereo view of a C^α trace of molecule B in the crystal of HSV I POL.

Figure 11 is an illustration of a ribbon diagram showing the overall structure of HSV 1 POL, and indicating the general location of the 3 -5' exonuclease active site (A), the extreme N-terminal domain (B), the N-terminal domain (C), the polymerase active site (D), the polymerase fingers domain (E), the polymerase palm domain (F), and the polymerase thumb domain (G).

Figure 12 illustrates a stereo view of the active site of the HSV 1 POL protein with PNU- 183792 and DNA duplex strands modeled. The residues listed in Tables 2-4 are shown in dark lines. Figure 13 is an illustration of the chemical structure of PNU- 183792 as disclosed, for example, in Thomsen et al., J. of Virology, 77:1868-76 (2003).

DETAILED DESCRIPTION OF PREFERRED EMBODIMENTS

Table 1 lists atomic structure coordinates derived by x-ray diffraction for crystals of Heφes simplex virus 1 DNA polymerase. Column 2 lists a number for the atom in the structure. Columns 3 and 12 list the element whose coordinates are measured (with the first letter(s) in column 3 defining the element). If the element is part of an amino acid, column 4 lists the type of amino acid using the three-letter abbreviations listed herein above; if the element is part of a water molecule, column 4 lists HOH; if the element is part of a sulfate anion, column 4 lists SO4; and if the element is part of a guanidine residue, column 4 lists GAI. Column 5 lists the chain id (A or B for each HSV 1 POL molecule in the asymmetric unit and W for solvent molecules, including water molecules, sulfate anions, and/or guanidine residues). Column 6 lists a number for the amino acid or molecule in the structure. Columns 7-9 list the crystallographic coordinates X, Y, and Z respectively. The crystallographic coordinates define the atomic position of the element measured. Column 10 lists an occupancy factor that refers to the fraction of the molecules in which each atom occupies the position specified by the coordinates. A value of "1" indicates that each atom has the same conformation, i.e., the same position, in all molecules of the crystal. Column 11 lists a thermal factor "B" that measures movement of the atom around its atomic center.

CONSTRUCTION OF FUSION PROTEIN A plasmid vector, pAcMAL (SEQ ID NO:5), was constructed to allow for simple cloning and isolation of a recombinant baculovirus that expresses foreign peptides in insect cells as fusion proteins. The desired peptide is fused to the C-terminus of the E. coli maltose binding protein (MBP). MBP is a periplasmic protein that binds to a cross-linked amylose resin. It has an amino-terminal signal peptide required for transport into the periplasm. The protein can be expressed intracellularly by removal of the DNA sequences encoding the signal peptide (Guan et al., Gene, 67:21-30 (1988)). Fusion proteins containing MBP preferably can bind to cross-linked amylose resin and can be specifically eluted from the column with maltose (Maina et al., Gene, 74:365- 373 (1988)). In addition, MBP portion of the fusion protein can be cleaved off by a protease.

A vector was constructed to use the strong baculovirus polyhedrin promoter to drive expression of the MBP-fusion protein (SEQ ID NO:3). The DNA polymerase gene from Heφes simplex virus type 1 (HSV-1) was cloned into pAcMAL and a recombinant baculovirus that expressed the entire HSV-1 DNA polymerase as an MBP-fusion protein was isolated. Immunoprecipitation experiments demonstrated that the fusion protein was mainly localized in the cytoplasm of infected insect cells.

It is well known that when expressed in E. coli, fusion partners may enhance target protein solubility and facilitate protein purification (Sambrook et al. in Molecular Cloning, Vol. 3, Chapter 15, Cold Spring Harbor Lab (2001)). However, there are few reports on fusion protein expression in baculovirus/insect cell system. A higher level of expression was observed with the fusion protein when compared to baculovirus expressing the non-fusion form of HSV-1 DNA polymerase. The concept of making a fusion protein to improve protein expression and purification may well be applicable to other members of Heφes DNA POL family. Indeed, high level of protein expression was also observed on MBP-Εpstein-Barr DNA polymerase fusion protein (MBP-ΕBV POL). Removing fusion partner from target protein can be achieved by endogeneous protease digestion as in the case of MBP-HSV 1 POL. However, to have better control of the digestion process, a second protease site was engineered by modifying the linker sequence between MBP and HSV 1 POL to introduce a second protease site (factor Xa in Figure 8). Removing of MBP at the second protease site is easily achieved without internal digestion. Because primary sequence analysis and crystal structure suggest that the first 42 residues of HSV 1 POL locate at the protein surface and are very flexible, they may serve as excellent linker at which controllable digestion can occur, in case of fusion proteins of other heφes DNA polymerase of which the N terminal peptides are not so flexible. For example, MBP-ΕBV POL fusion protein digestion is quite difficult without internal digestion due to lack of flexible linker, but the linker from MBP-HSV 1 POL may render it much easier. An exposed His tag may also be introduced in this linker region, facilitating possible affinity purification by immobilized metal resin.

Expression conditions for heφes simplex virus 1 polymerase (HSV 1 POL) were then optimized, and high level protein expression was achieved in the baculovirus/insect cell system in a bioreactor. The expressed HSV 1 POL fusion protein (SEQ ID NO: 3) included MBP fused to the N terminus of the polymerase. After cell disruption, MBP-HSV 1 POL was extracted with high salt. The extracted protein was captured by a cation exchange column such as phosphocellulose column. The release of HSV 1 POL (SEQ ID NO: 1) from the fusion protein was realized with digestions by either endogenous proteases of the expression system or by a specific protease such as factor Xa on the modified construct with two protease cleavage sites (factor Xa in Figure 8). The product was separated from the undigested full-length fusion protein and some other impurities on a hydroxyapatite (HA) column. Additional minor impurities were removed on a affinity column, such as a DNA-cellulose column and a heparin column (Figure 9). Homogeneity of the purified protein was verified by SDS- PAGE gel, N-terminal sequencing, amino acid composition analysis, and mass spectroscopy. HSV 1 POL activity was confirmed by isotope labeled nucleotide incoiporation assays.

CRYSTALLINE FORM(S) AND METHOD OF MAKING

The three-dimensional structure of Heφes simplex virus 1 DNA polymerase was solved using x-ray crystallography to 2.8 A resolution for a Heφes simplex virus 1 DNA polymerase crystal.

Preferably, the crystal has orthorhombic space group symmetry ^~?2{2{2_\. Preferably, the crystal includes parallelepiped shaped unit cells, each unit cell having dimensions a=105 ± 25 A, b=125 ± 25 A, c=225 + 25 A, and α=β=- 90°. More preferably, the crystal includes parallelepiped shaped unit cells, each unit cell having dimensions a=103 ± 5 A, b=125 ± 5 A, c=222 ± 5 A, and =β=γ=90°. Most preferably, the crystal includes parallelepiped shaped unit cells, each unit cell having dimensions of about a=103 A, b=125 A, c=222 A, and α=β=γ=90°. Preferably, the crystallized enzyme is a monomer having two monomers in the crystallographic asymmetric unit. For crystallization, preferably, HSV 1 POL is concentrated to 25+24 mg/ml in a buffer solution of 100+80 mM HEPES, pH=7.5±2.5, 20+10% glycerol, 5+4mM DTT, 400+.200 mM guanidine-HCl and 1±0.9 mM EDTA. More preferably, HSV 1 POL is concentrated to 25+10 mg/ml in a buffer solution of 60+40 mM HEPES, pH=7.5±l .5, 15+5% glycerol, 3+2mM DTT, 200+100 mM guanidine-HCl and 0.6+0.5 mM EDTA. Most preferably, HSV 1 POL is concentrated to 25 mg/ml in a buffer solution of 20 mM HEPES, pH=7.5, 10% glycerol, 5mM DTT, 200+100 mM guanidine and ImM EDTA. Preferably, a method of slow exchange of protein buffer with precipitant solution is used for crystallization of HSV 1 POL. More preferably, a vapor diffusion against precipitant solution method is used for crystallization of HSV 1 POL. Most preferably, a hanging drop vapor diffusion against precipitant solution method is used for crystallization of HSV 1 POL. Preferably, the precipitant solution is of 200+190 mM HEPES, pH 7.0+2.5, 200+190 mM ammonium sulfate, 10+9 mM DTT, 200+190 mM guanidine-HCl, 0-25% of PEG-1000, PEG-2000, PEG-3350, PEG-4000, PEG-6000, PEG-8000, PEG- 10,000, or PEG-20,000. More preferably, the precipitant solution is of 150+100 mM HEPES, pH 7.0+1.5, 100+90 mM ammonium sulfate, 5+4 mM DTT, 150+100 mM guanidine-HCl, 0- 15% of PEG-2000, PEG-3350, PEG-4000, PEG-6000 or PEG-8000. Most preferably, the precipitant solution is of 100 mM HEPES, pH 7.0, 50 mM ammonium sulfate, 5 mM DTT, 100 mM guanidine-HCl, 4% PEG-3350. Preferably, before crystallization organic cosmotropic solutes such as DMSO, or glycerol, or ethyleneglycol, or xylitol, or sorbitol, or glucose, or sucrose, or trehalose, or β-alanine or sarcosine are added into the protein solution to a concentration of 1-30%. More preferably, DMSO is added to a concentration of 5-15%. Most preferably, DMSO is added to a concentration of 11.1%. In order to collect complete data on single Heφes simplex virus 1 DNA polymerase crystals, crystals have to exchanged into a cryo-protection solution before freezing in liquid nitrogen. Preferably, the cryo-protection solution includes: 20- 200 mM HEPES, pH 5-10, 20-120 mM ammonium sulfate, 1-10 mM DTT, 10- 400 mM guanidine-HCl, 1-30% PEG-3350, 1-30% DMSO, 1-30% PEG-400, or PEG-600, or PEG-1000. More preferably, the cryo-protection solution includes: 50-150 mM HEPES, pH 6.5-8.5, 40-80 mM ammonium sulfate, 3-8 mM DTT, 50-150 mM guanidine-HCl, 5-15% PEG-3350, 5-15% DMSO, 5-15% PEG- 1000. Most preferably, the cryo-protection solution includes: 100 mM HEPES, pH 7, 60 mM ammonium sulfate, 5 mM DTT, 100 mM guanidine-HCl, 10% PEG-3350, 11% DMSO, 10% PEG-1000.

Variation in buffer and buffer pH as well as other additives such as PEG, PEG-MME, PEG-DME, or polyoxyalkylenepolyamines is apparent to those skilled in the art and may result in similar crystals.

The invention further includes a Heφes simplex virus 1 DNA polymerase crystal that is isomoφhous with a Heφes simplex virus 1 DNA polymerase crystal having a unit cell defined by the dimensions of a, b, c, , β, and γ, wherein a is about 80 A to about 130 A, b is about 100 A to about 150 A, c is about 200 A to about 250 A, and α=β=Υ=90°.

Crystallography may preferably be used to screen and identify chemical entities that are not known potential modifiers of target biomolecules as disclosed, for example, in U.S. Pat. No. 6,297,021 (Nienaber et al.). As used herein, a "potential modifier" refers to a chemical entity that could function as a drug candidate (e.g., modifiers and inhibitors). For example, crystallography may preferably be used to screen and identify chemical entities that are not known potential modifiers of Heφes simplex virus 1 DNA polymerase for their ability to bind to Heφes simplex virus 1 DNA polymerase. A preferred method includes obtaining a crystal of Heφes simplex virus 1 DNA polymerase; exposing the Heφes simplex virus 1 DNA polymerase to one or more test samples that include a potential modifier of the Heφes simplex virus 1 DNA polymerase; and determining whether a potential modifier-Heφes simplex virus 1 DNA polymerase molecular complex is formed. The Heφes simplex virus 1 DNA polymerase may be exposed to potential modifiers by various methods including, for example, soaking a Heφes simplex virus 1 DNA polymerase crystal in a solution of one or more potential modifiers, or co-crystallizing Heφes simplex virus 1 DNA polymerase in the presence of one or more potential modifiers. Structural information from the potential modifier-Heφes simplex virus

1 DNA polymerase complexes found may preferably be used to design new potential modifiers that bind tighter, bind more specifically, have desired biological activity properties, have better safety profiles than known potential modifiers, and combinations thereof. For example, libraries of "shape-diverse" chemical entities may preferably be used to allow direct identification of the potential modifier-Heφes simplex virus 1 DNA polymerase complex even when the potential modifier is exposed as part of a mixture. As used herein, "shape diverse" refers to potential modifiers having substantial differences in three- dimensional shapes that can be recognized, for example, by visual inspection of the two dimensional chemical structures, or by calculation and comparison of relevant parameters by a computational program. Shape diversity of the mixture permits a bound potential modifier to be identified directly from the resultant electron density map. This preferably avoids the need for time-consuming deconvolution of a hit from the mixture. Here, three important steps are preferably achieved simultaneously. Preferably, the calculated electron density function directly reveals the binding event, identifies the bound chemical entity, and provides a detailed 3-D structure of the potential modifier-Heφes simplex virus 1 DNA polymerase complex. Once a hit is found, preferably a number of analogs or derivatives of the hit may be screened for tighter binding or desired biological activity by traditional screening methods. Moreover, the identity of the hit and information about structure of the target may preferably be used to develop analogs or derivatives with tighter binding or desired biological activity properties. Optionally, the potential modifier-Heφes simplex virus 1 DNA polymerase complex may be exposed to additional iterations of potential modifiers so that two or more hits may preferably be linked together to identify or design a more potent potential modifier.

X-RAY CRYSTALLOGRAPHIC ANALYSIS Each of the constituent amino acids of Heφes simplex virus 1 DNA polymerase is defined by a set of structure coordinates as set forth in Table 1. The term "structure coordinates" refers to Cartesian coordinates derived from mathematical equations related to the patterns obtained on diffraction of a monochromatic beam of x-rays by the atoms (scattering centers) of a Heφes simplex virus 1 DNA polymerase complex in crystal form. The diffraction data are used to calculate an electron density map of the repeating unit of the crystal. The electron density maps are then used to establish the positions of the individual atoms of the Heφes simplex virus 1 DNA polymerase protein or protein-potential modifier complex. Slight variations in structure coordinates can be generated by mathematically manipulating the Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-potential modifier structure coordinates. For example, the structure coordinates set forth in Table 1 could be manipulated by crystallographic permutations of the structure coordinates, fractionalization of the structure coordinates, integer additions or subtractions to sets of the structure coordinates, inversion of the structure coordinates or any combination of the above. Alternatively, modifications in the crystal structure due to mutations, additions, substitutions, and/or deletions of amino acids, or other changes in any of the components that make up the crystal, could also yield variations in structure coordinates. Such slight variations in the individual coordinates will have little effect on overall shape. If such variations are within an acceptable standard error as compared to the original coordinates, the resulting three-dimensional shape is considered to be structurally equivalent. Structural equivalence is described in more detail below. It should be noted that slight variations in individual structure coordinates of the Heφes simplex virus 1 DNA polymerase would not be expected to significantly alter the nature of chemical entities such as potential modifiers that could associate with the active sites. In this context, the phrase "associating with" refers to a condition of proximity between a chemical entity, or portions thereof, and a Heφes simplex virus 1 DNA polymerase molecule or portions thereof. The association may be non-covalent, wherein the juxtaposition is energetically favored by hydrogen bonding, van der Waals forces, or electrostatic interactions, or it may be covalent. Thus, for example, a potential modifier that bound to an active site of Heφes simplex virus 1 DNA polymerase would also be expected to bind to or interfere with a structurally equivalent active site.

For the puφose of this invention, any molecule or molecular complex or active site thereof, or any portion thereof, that has a root mean square deviation of conserved residue backbone atoms (N, Cα, C, O) of less than about 1 A, when superimposed on the relevant backbone atoms described by the reference structure coordinates listed in Table 1, is considered "structurally equivalent" to the reference molecule. That is to say, the crystal structures of those portions of the two molecules are substantially identical, within acceptable error. As used herein, "residue" refers to one or more atoms. Particularly preferred structurally equivalent molecules or molecular complexes are those that are defined by the entire set of structure coordinates listed in Table 1 ± a root mean square deviation from the conserved backbone atoms of those amino acids of less than about 1 A. More preferably, the root mean square deviation is at most about 0.5 A, and even more preferably, at most about 0.35 A. Other embodiments of this invention include a molecular complex defined by the structure coordinates listed in Table 1 for those amino acids listed in Table 2, Table 3, or Table 4 ± a root mean square deviation from the conserved backbone atoms of those amino acids of less than about 3 A, preferably at most about 2 A, and more preferably at most about 1 A. The term "root mean square deviation" means the square root of the arithmetic mean of the squares of the deviations. It is a way to express the deviation or variation from a trend or object. For puφoses of this invention, the "root mean square deviation" defines the variation in the backbone of a protein from the backbone of Heφes simplex virus 1 DNA polymerase or an active site portion thereof, as defined by the structure coordinates of Heφes simplex virus 1 DNA polymerase described herein.

It will be readily apparent to those of skill in the art that the numbering of amino acids in other isoforms of Heφes simplex virus 1 DNA polymerase may be different than that of Heφes simplex virus 1 DNA polymerase used in this application.

OVERVIEW OF HSV I POL STRUCTURES

There are two independent HSV 1 POL molecules in the crystallographic asymmetric unit (Figure 10). The overall structures are very similar between these two molecules but local variations do occur. Roughly, the molecules can be divided into domains as illustrated in Figure 11 : the extreme N-terminal domain (e.g., from the N-terminus to residue 140; "B"); the N-terminal domain (e.g., residues 141-353 and 593-639; "C"); the 3'-5' exonuclease domain (e.g., residues 362-592; "A"); the polymerase palm domain (e.g., residues 701-766 and 824-955; "F"); the polymerase fingers domain (e.g., residues 767-825; "E"); the polymerase thumb domain (e.g., residues 963-1197; "G"); and the accessory protein binding domain (e.g., residues 1200-1235; not ordered in this structure and not illustrated in Figure 11). The extreme N-terminal domain is located at the outer surface of the 3 '-5' exonuclease domain and is very loosely packed with only a few secondary structures. With the truncated N-terminal 42 amino acids of the protein, the first amino acid residue visible in the electron density map is residue 59. The N-terminal domain consists of two β-sheet and contacts the fingers through its last portion of two -helices. The 3'-5' exo-nuclease domain consists a central β-sheet, flanked by 6 α-helices. The polymerase domain is sub-divided into palm, fingers, and thumb because the domain itself reassembles a right hand, with the palm domain in the middle. The whole molecule forms a circular disk structure, with the thumb sub-domain of the polymerase touching the 3 '-5' exonuclease domain. The overall structure is quite similar to that of the bacterial phage RB69

DNA polymerase (Wang et al., Cell, 89:1087-1099 (1997); Shamoo et al., Cell, 99:155-166 (1999); and Franklin et al., Cell, 105:657-667 (2001)). The topology of 3 '-5' exonuclease and polymerase domain are the same as RB69. However, large differences are also observed, especially on their N-terminal domains and polymerase thumb subdomains. Relative domain movements are also observed due to conformational changes. Positions of the fingers and the 3'-5' exonuclease domain of HSV 1 POL relatively to the palm domain are closer to those observed in apo RB69 structure. RMS difference is 0.74A when peptides 407-413, 587-601 and 624-629 of phage DNA polymerase palm domain superimposed with 713-719, 838-852 and 889-894 of HSV 1 POL.

ACTIVE SITE AND OTHER STRUCTURAL FEATURES

Applicants' invention provides information about the shape and structure of the active sites of Heφes simplex virus 1 DNA polymerase in the presence of a potential modifier.

Active sites are of significant utility in fields such as drug discovery. The association of natural potential modifiers or substrates with the active sites of their corresponding receptors or enzymes is the basis of many biological mechanisms of action. Similarly, many drugs exert their biological effects through association with the active sites of receptors and enzymes. Such associations may occur with all or any parts of the active site. An understanding of such associations helps lead to the design of drugs having more favorable associations with their target, and thus improved biological effects. Therefore, this information is valuable in designing potential modifiers of Heφes simplex virus 1 DNA polymerase-like active sites, as discussed in more detail below. The term "active site," as used herein, refers to a region of a molecule or molecular complex, that, as a result of its shape, favorably associates with another chemical entity. Thus, an active site may include or consist of features such as cavities, surfaces, or interfaces between domains. Chemical entities that may associate with an active site include, but are not limited to, cofactors, substrates, modifiers, agonists, and antagonists.

The amino acid constituents of a Heφes simplex virus 1 DNA polymerase active site as defined herein are positioned in three dimensions in accordance with the structure coordinates listed in Table 1. In one aspect, the structure coordinates defining an active site of Heφes simplex virus 1 DNA polymerase include structure coordinates of all atoms in the constituent amino acids; in another aspect, the structure coordinates of an active site include structure coordinates of just the backbone atoms of the constituent amino acids. Heφes simplex virus 1 DNA polymerase has at least two distinct and well defined active sites: polymerase active site and 3 '-5' exonuclease active site. In one embodiment, the polymerase active site of Heφes simplex virus 1 DNA polymerase preferably includes the amino acids listed in Table 2, more preferably the amino acids listed in Table 3, and most preferably the amino acids listed in Table 4, as represented by the structure coordinates listed in Table 1. The polymerase active site includes nucleotide residues in addition to the amino acid residues described herein above. The nucleotide residues can be identified by methods known to those of skill in the art including, for example, the modeling exercise described herein above. Alternatively, the polymerase active site of Heφes simplex virus 1 DNA polymerase may be defined by those amino acids and nucleotides whose backbone atoms are situated within about 4 A, more preferably within about 7 A, most preferably within about 10 A, of one or more constituent atoms of a bound substrate or modifier. In yet another alternative, the polymerase active site may be defined by those amino acids whose backbone atoms are situated within a sphere centered on the coordinates representing the alpha carbon atom of residue Asp888, the sphere having a radius of about 25 A, preferably about 15 A, and more preferably about 10 A.

In another embodiment, the 3'-5' exonuclease active site of Heφes simplex virus 1 DNA polymerase preferably includes the amino acids listed in Table 5, more preferably the amino acids listed in Table 6, and most preferably the amino acids listed in Table 7, as represented by the structure coordinates listed in Table 1. Alternatively, the 3 -5' exonuclease active site of Heφes simplex virus 1 DNA polymerase may be defined by those amino acids and nucleotides whose backbone atoms are situated within about 4 A, more preferably within about 7 A, most preferably within about 10 A, of one or more constituent atoms of a bound substrate or modifier. In yet another alternative, the 3'-5' exonuclease active site may be defined, for example, by those amino acids whose backbone atoms are situated within a sphere centered on the coordinates representing the alpha carbon atom of residue Glu370, the sphere having a radius of about 25 A, preferably about 15 A, and more preferably about 10 A.

The term "Heφes simplex virus 1 DNA polymerase-like active site" refers to a portion of a molecule or molecular complex whose shape is sufficiently similar to at least a portion of an active site of Heφes simplex virus 1 DNA polymerase as to be expected to bind related structural analogues. As used herein, "at least a portion" means that at least about 50% of the amino acids are included, preferably at least about 70% of the amino acids are included, more preferably at least about 90% of the amino acids are included, and most preferably all the amino acids are included. A structurally equivalent active site is defined by a root mean square deviation from the structure coordinates of the backbone atoms of the amino acids that make up active sites in Heφes simplex virus 1 DNA polymerase (as set forth in Table 1) of at most about 1 A. How this calculation is obtained is described below.

Accordingly, the invention provides molecules or molecular complexes including a Heφes simplex virus 1 DNA polymerase active site or Heφes simplex virus 1 DNA polymerase-like active site, as defined by the sets of structure coordinates described above.

TABLE 2: Residues with 4A of Polymerase Active Site.

TABLE 4: Residues with lOA of Polymerse Active Site.

TABLE 5: Residues with 4A of 3 -5' Exonuclease Active Site.

TABLE 6: Residues with 7A of 3'-5' Exonuclease Active Site.

TABLE 7: Residues with 10A of 3 -5' Exonuclease Active Site.

THREE-DIMENSIONAL CONFIGURATIONS

X-ray structure coordinates define a unique configuration of points in space. Those of skill in the art understand that a set of structure coordinates for protein or an protein-potential modifier complex, or a portion thereof, define a relative set of points that, in turn, define a configuration in three dimensions. A similar or identical configuration can be defined by an entirely different set of coordinates, provided the distances and angles between coordinates remain essentially the same. In addition, a scalable configuration of points can be defined by increasing or decreasing the distances between coordinates by a scalar factor while keeping the angles essentially the same. The present invention thus includes the scalable three-dimensional configuration of points derived from the structure coordinates of at least a portion of a Heφes simplex virus 1 DNA polymerase molecule or molecular complex, as listed in Table 1, as well as structurally equivalent configurations, as described below. Preferably, the scalable three-dimensional configuration includes points derived from structure coordinates representing the locations of a plurality of the amino acids defining a Heφes simplex virus 1 DNA polymerase active site.

In one embodiment, the scalable three-dimensional configuration includes point^'s derived from structure coordinates representing the locations the backbone atoms of a plurality of amino acids defining the Heφes simplex virus 1 DNA polymerase active site, preferably the amino acids listed in Table 2, more preferably the amino acids listed in Table 3, and most preferably the amino acids listed in Table 4.

Alternatively, the scalable three-dimensional configuration includes points derived from structure coordinates representing the locations of the side chain and the backbone atoms (other than hydrogens) of a plurality of the amino acids defining the Heφes simplex virus 1 DNA polymerase active site, preferably the amino acids listed in Table 2, more preferably the amino acids listed in Table 3, and most preferably the amino acids listed in Table 4. Likewise, the invention also includes the scalable three-dimensional configuration of points derived from structure coordinates of molecules or molecular complexes that are structurally homologous to Heφes simplex virus 1 DNA polymerase, as well as structurally equivalent configurations. Structurally homologous molecules or molecular complexes are defined below. Advantageously, structurally homologous molecules can be identified using the structure coordinates of Heφes simplex virus 1 DNA polymerase according to a method of the invention.

The configurations of points in space derived from structure coordinates according to the invention can be visualized as, for example, a holographic image, a stereodiagram, a model, or a computer-displayed image, and the invention thus includes such images, diagrams or models.

STRUCTURALLY EQUIVALENT CRYSTAL STRUCTURES

Various computational analyses can be used to determine whether a molecule or an active site portion thereof is "structurally equivalent," defined in terms of its three-dimensional structure, to all or part of Heφes simplex virus 1 DNA polymerase or its active sites. Such analyses may be carried out in current software applications, such as the Molecular Similarity application of QUANTA (Molecular Simulations Inc., San Diego, CA) version 4.1, and as described in the accompanying User's Guide.

The Molecular Similarity application permits comparisons between different structures, different conformations of the same structure, and different parts of the same structure. The procedure used in Molecular Similarity to compare structures is divided into four steps: (1) load the structures to be compared; (2) define the atom equivalences in these structures; (3) perform a fitting operation; and (4) analyze the results.

Each structure is identified by a name. One structure is identified as the target (i.e., the fixed structure); all remaining structures are working structures (i.e., moving structures). Since atom equivalency within QUANTA is defined by user input, for the puφose of this invention equivalent atoms are defined as protein backbone atoms (N, Cα, C, and O) for all conserved residues between the two structures being compared. A conserved residue is defined as a residue which is structurally or functionally equivalent. Only rigid fitting operations are considered. When a rigid fitting method is used, the working structure is translated and rotated to obtain an optimum fit with the target structure. The fitting operation uses an algorithm that computes the optimum translation and rotation to be applied to the moving structure, such that the root mean square difference of the fit over the specified pairs of equivalent atom is an absolute minimum. This number, given in angstroms, is reported by QUANTA.

MACHINE READABLE STORAGE MEDIA

Transformation of the structure coordinates for all or a portion of Heipes simplex virus 1 DNA polymerase or the Heφes simplex virus 1 DNA polymerase-potential modifier complex or one of its active sites, for structurally homologous molecules as defined below, or for the structural equivalents of any of these molecules or molecular complexes as defined above, into three- dimensional graphical representations of the molecule or complex can be conveniently achieved through the use of commercially-available software.

The invention thus further provides a machine-readable storage medium including a data storage material encoded with machine readable data which, when using a machine programmed with instructions for using said data, displays a graphical three-dimensional representation of any of the molecule or molecular complexes of this invention that have been described above. In a preferred embodiment, the machine-readable data storage medium includes a data storage material encoded with machine readable data which, when using a machine programmed with instructions for using said data, displays a graphical three- dimensional representation of a molecule or molecular complex including all or any parts of a Heφes simplex virus 1 DNA polymerase active site or an Heφes simplex virus 1 DNA polymerase-like active site, as defined above. In another preferred embodiment, the machine-readable data storage medium includes a data storage material encoded with machine readable data which, when using a machine programmed with instructions for using said data, displays a graphical three-dimensional representation of a molecule or molecular complex defined by the structure coordinates of all of the amino acids listed in Table 1, ± a root mean square deviation from the backbone atoms of said amino acids of less than about 0.365 A, more preferably at most about 2 A, and even more preferably, at most about 1 A. In an alternative embodiment, the machine-readable data storage medium includes a data storage material encoded with a first set of machine readable data which includes the Fourier transform of the structure coordinates set forth in Table 1, and which, when using a machine programmed with instructions for using said data, can be combined with a second set of machine readable data including the x-ray diffraction pattern of a molecule or molecular complex to determine at least a portion of the structure coordinates corresponding to the second set of machine readable data.

For example, a system for reading a data storage medium may include a computer including a central processing unit ("CPU"), a working memory which may be, e.g., RAM (random access memory) or "core" memory, mass storage memory (such as one or more disk drives or CD-ROM drives), one or more display devices (e.g., cathode-ray tube ("CRT") displays, light emitting diode ("LED") displays, liquid crystal displays ("LCDs"), electroluminescent displays, vacuum fluorescent displays, field emission displays ("FEDs"), plasma displays, projection panels, etc.), one or more user input devices (e.g., keyboards, microphones, mice, track balls, touch pads, etc.), one or more input lines, and one or more output lines, all of which are interconnected by a conventional bidirectional system bus. The system may be a stand-alone computer, or may be networked (e.g., through local area networks, wide area networks, intranets, extranets, or the internet) to other systems (e.g., computers, hosts, servers, etc.). The system may also include additional computer controlled devices such as consumer electronics and appliances.

Input hardware may be coupled to the computer by input lines and may be implemented in a variety of ways. Machine-readable data of this invention may be inputted, for example, via the use of a modem or modems connected by a telephone line or dedicated data line or via satellite down loads. Alternatively or additionally, the input hardware may include CD-ROM drives or disk drives. In conjunction with a display terminal, a keyboard may also be used as an input device. Output hardware may be coupled to the computer by output lines and may similarly be implemented by conventional devices. By way of example, the output hardware may include a display device for displaying a graphical representation of an active site of this invention using a program such as QUANTA as described herein. Output hardware might also include a printer, so that hard copy output may be produced, or a disk drive, to store system output for later use.

In operation, a CPU coordinates the use of the various input and output devices, coordinates data accesses from mass storage devices, accesses to and from working memory, and determines the sequence of data processing steps. A number of programs may be used to process the machine-readable data of this invention. Such programs are discussed in reference to the computational methods of drug discovery as described herein. References to components of the hardware system are included as appropriate throughout the following description of the data storage medium. Machine-readable storage devices useful in the present invention include, but are not limited to, magnetic devices, electrical devices, optical devices, and combinations thereof. Examples of such data storage devices include, but are not limited to, hard disk devices, CD devices, digital video disk devices, floppy disk devices, removable hard disk devices, magneto-optic disk devices, magnetic tape devices, flash memory devices, bubble memory devices, holographic storage devices, and any other mass storage peripheral device. It should be understood that these storage devices include necessary hardware (e.g., drives, controllers, power supplies, etc.) as well as any necessary media (e.g., disks, flash cards, etc.) to enable the storage of data. STRUCTURALLY HOMOLOGOUS MOLECULES, MOLECULAR COMPLEXES, AND CRYSTAL STRUCTURES

The structure coordinates set forth in Table 1 can be used to aid in obtaining structural information about another crystallized molecule or molecular complex. The method of the invention allows determination of at least a portion of the three-dimensional structure of molecules or molecular complexes which contain one or more structural features that are similar to structural features of Heφes simplex virus 1 DNA polymerase. These molecules are referred to herein as "structurally homologous" to Heφes simplex virus 1 DNA polymerase. Similar structural features can include, for example, regions of amino acid identity, conserved active site or active site motifs, and similarly arranged secondary structural elements (e.g., α helices and β sheets). Optionally, structural homology is determined by aligning the residues of the two amino acid sequences to optimize the number of identical amino acids along the lengths of their sequences; gaps in either or both sequences are permitted in making the alignment in order to optimize the number of identical amino acids, although the amino acids in each sequence must nonetheless remain in their proper order. Preferably, two amino acid sequences are compared using the Blastp program, version 2.0.9, of the BLAST 2 search algorithm, as described by Tatusova et al., FEMS Microbiol Lett., 174:247-50 (1999), and available on the world wide web at ncbi.nlm.nih.gov/gorf/bl2.html. Preferably, the default values for all BLAST 2 search parameters are used, including matrix = BLOSUM62; open gap penalty = 11, extension gap penalty = 1, gap x_dropoff = 50, expect = 10, wordsize = 3, and filter on. In the comparison of two amino acid sequences using the BLAST search algorithm, structural similarity is referred to as "identity." Preferably, a structurally homologous molecule is a protein that has an amino acid sequence sharing at least 65% identity with a native or recombinant amino acid sequence of Heφes simplex virus 1 DNA polymerase (for example, SEQ ID NO:l). More preferably, a protein that is structurally homologous to Heφes simplex virus 1 DNA polymerase includes a contiguous stretch of at least 50 amino acids that shares at least 80% amino acid sequence identity with the analogous portion of the native or recombinant Heφes simplex virus 1 DNA polymerase (for example, SEQ ID NO:l). Methods for generating structural information about the structurally homologous molecule or molecular complex are well-known and include, for example, molecular replacement techniques.

Therefore, in another embodiment this invention provides a method of utilizing molecular replacement to obtain structural information about a molecule or molecular complex whose structure is unknown including the steps of: (a) crystallizing the molecule or molecular complex of unknown structure;

(b) generating an x-ray diffraction pattern from said crystallized molecule or molecular complex; and

(c) applying at least a portion of the structure coordinates set forth in Table 1 to the x-ray diffraction pattern to generate a three-dimensional electron density map of the molecule or molecular complex whose structure is unknown.

By using molecular replacement, all or part of the structure coordinates of Heφes simplex virus 1 DNA polymerase or the Heφes simplex virus 1 DNA polymerase-potential modifier complex as provided by this invention can be used to determine the structure of a crystallized molecule or molecular complex whose structure is unknown more quickly and efficiently than attempting to determine such information ab initio.

Molecular replacement provides an accurate estimation of the phases for an unknown structure. Phases are a factor in equations used to solve crystal structures that cannot be determined directly. Obtaining accurate values for the phases, by methods other than molecular replacement, is a time-consuming process that involves iterative cycles of approximations and refinements and greatly hinders the solution of crystal structures. However, when the crystal structure of a protein containing at least a structurally homologous portion has been solved, the phases from the known structure provide a satisfactory estimate of the phases for the unknown structure.

Thus, this method involves generating a preliminary model of a molecule or molecular complex whose structure coordinates are unknown, by orienting and positioning the relevant portion of Heφes simplex virus 1 DNA polymerase or the Heφes simplex virus 1 DNA polymerase/modifier complex within the unit cell of the crystal of the unknown molecule or molecular complex so as best to account for the observed x-ray diffraction pattern of the crystal of the molecule or molecular complex whose structure is unknown. Phases can then be calculated from this model and combined with the observed x-ray diffraction pattern amplitudes to generate an electron density map of the structure whose coordinates are unknown. This, in turn, can be subjected to any well-known model building and structure refinement techniques to provide a final, accurate structure of the unknown crystallized molecule or molecular complex (Lattman, Meth. EnzymoL, 115, 55-77 (1985); M.G. Rossman, ed., "The Molecular

Replacement Method," Int. Sci. Rev. Ser., No. 13, Gordon & Breach, New York (1972)).

Structural information about a portion of any crystallized molecule or molecular complex that is sufficiently structurally homologous to a portion of Heφes simplex virus 1 DNA polymerase can be resolved by this method. In addition to a molecule that shares one or more structural features with Heφes simplex virus 1 DNA polymerase as described above, a molecule that has similar bioactivity, such as the same catalytic activity, substrate specificity, or potential modifier binding activity as Heφes simplex virus 1 DNA polymerase, may also be sufficiently structurally homologous to Heφes simplex virus 1 DNA polymerase to permit use of the structure coordinates of Heφes simplex virus 1 DNA polymerase to solve its crystal structure.

In a preferred embodiment, the method of molecular replacement is utilized to obtain structural information about a molecule or molecular complex, wherein the molecule or molecular complex includes a Heφes simplex virus 1 DNA polymerase subunit or homolog. A "subunit" of Heφes simplex virus 1 DNA polymerase is a Heφes simplex virus 1 DNA polymerase molecule that has been truncated at the N-terminus or the C-terminus, or both. In the context of the present invention, a "homolog" of Heφes simplex virus 1 DNA polymerase is a protein that contains one or more amino acid substitutions, deletions, additions, or rearrangements with respect to the amino acid sequence of Heφes simplex virus 1 DNA polymerase (SEQ ID NO:l), but that, when folded into its native conformation, exhibits or is reasonably expected to exhibit at least a portion of the tertiary (three-dimensional) structure of Heφes simplex virus 1 DNA polymerase. For example, structurally homologous molecules can contain deletions or additions of one or more contiguous or noncontiguous amino acids, such as a loop or a domain. Structurally homologous molecules also include "modified" Heφes simplex virus 1 DNA polymerase molecules that have been chemically or enzymatically derivatized at one or more constituent amino acid, including side chain modifications, backbone modifications, and N- and C- terminal modifications including acetylation, hydroxylation, methylation, amidation, and the attachment of carbohydrate or lipid moieties, cofactors, and the like.

A heavy atom derivative of Heφes simplex virus 1 DNA polymerase is also included as a Heφes simplex virus 1 DNA polymerase homolog. The term "heavy atom derivative" refers to derivatives of Heφes simplex virus 1 DNA polymerase produced by chemically modifying a crystal of Heφes simplex virus 1 DNA polymerase. In practice, a crystal is soaked in a solution containing heavy metal atom salts, or organometallic compounds, e.g., lead chloride, gold thiomalate, thiomersal or uranyl acetate, which can diffuse through the crystal and bind to the surface of the protein. The location(s) of the bound heavy metal atom(s) can be determined by x-ray diffraction analysis of the soaked crystal. This information, in turn, is used to generate the phase information used to construct three-dimensional structure of the protein (T.L. Blundell and N.L. Johnson, Protein Crystallography, Academic Press (1976)). Because Heφes simplex virus 1 DNA polymerase can crystallize in more than one crystal form, the structure coordinates of Heφes simplex virus 1 DNA polymerase as provided by this invention are particularly useful in solving the structure of other crystal forms of Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase complexes.

The structure coordinates of Heφes simplex virus 1 DNA polymerase as provided by this invention are particularly useful in solving the structure of Heφes simplex virus 1 DNA polymerase mutants. Mutants may be prepared, for example, by expression of Heφes simplex virus 1 DNA polymerase cDNA previously altered in its coding sequence by oligonucleotide-directed mutagenesis. Mutants may also be generated by site-specific incoφoration of unnatural amino acids into Heφes simplex virus 1 DNA polymerase proteins using the general biosynthetic method of Noren et al., Science, 244:182-88 (1989). In this method, the codon encoding the amino acid of interest in wild- type Heφes simplex virus 1 DNA polymerase is replaced by a "blank" nonsense codon, TAG, using oligonucleotide-directed mutagenesis. A suppressor tRNA directed against this codon is then chemically aminoacylated in vitro with the desired unnatural amino acid. The aminoacylated tRNA is then added to an in vitro translation system to yield a mutant Heφes simplex virus 1 DNA polymerase with the site-specific incoφorated unnatural amino acid.

Selenocysteine or selenomethionine may be incoφorated into wild-type or mutant Heφes simplex virus 1 DNA polymerase by expression of Heφes simplex virus 1 DNA polymerase-encoding cDNAs in baculovirus/insect cell system. In this method, the wild-type or mutagenized Heφes simplex virus 1 DNA polymerase cDNA may be expressed in a host organism on a growth medium depleted of either natural cysteine or methionine (or both) but enriched in selenocysteine or selenomethionine (or both) (Bellizzi HI et al., Proceedings of the National Academy of Sciences, U.S., 97:4573-4578 (2000)) . Alternatively one may use other methionine replacements (telluromethionine) that are useful for phasing. (Karnbrock et al., J. of Molecular Biology, 270:616-23 (1997).). The structure coordinates of Heφes simplex virus 1 DNA polymerase listed in Table 1 are also particularly useful to solve the structure of crystals of Heφes simplex virus 1 DNA polymerase, Heφes simplex virus 1 DNA polymerase mutants or Heφes simplex virus 1 DNA polymerase homologs co- complexed with a variety of chemical entities. This approach enables the determination of the optimal sites for interaction between chemical entities, including candidate Heφes simplex virus 1 DNA polymerase modifiers and Heφes simplex virus 1 DNA polymerase.

Potential sites for modification within the various active sites of the molecule can also be identified. This information provides an additional tool for determining the most efficient binding interactions, for example, increased hydrophobic interactions, between Heφes simplex virus 1 DNA polymerase and a chemical entity. For example, high resolution x-ray diffraction data collected from crystals exposed to different types of solvent allows the determination of where each type of solvent molecule resides. Small molecules that bind tightly to those sites can then be designed and synthesized and tested for their potential Heφes simplex virus 1 DNA polymerase inhibition activity.

All of the complexes referred to above may be studied using well-known x-ray diffraction techniques and may be refined versus 1.5-3.5 A resolution x-ray data to an R value of about 0.30 or less using computer software, such as X- PLOR (Yale University, 1992, distributed by Molecular Simulations, Inc.; see, e.g., Blundell & Johnson, supra; Meth. Enzymol, Vol. 114 & 115, H.W. Wyckoff et al., eds., Academic Press (1985)). This information may thus be used to optimize known Heφes simplex virus 1 DNA polymerase modifiers, and more importantly, to design new Heφes simplex virus 1 DNA polymerase modifiers.

The invention also includes the unique three-dimensional configuration defined by a set of points defined by the structure coordinates for a molecule or molecular complex structurally homologous to Heφes simplex virus 1 DNA polymerase as determined using the method of the present invention, structurally equivalent configurations, and magnetic storage media including such set of structure coordinates.

Further, the invention includes structurally homologous molecules as identified using the method of the invention.

HOMOLOGY MODELING

Using homology modeling, a computer model of a Heφes simplex virus 1 DNA polymerase homolog can be built or refined without crystallizing the homolog. First, a preliminary model of the Heφes simplex virus 1 DNA polymerase homolog is created by sequence alignment with Heφes simplex virus 1 DNA polymerase, secondary structure prediction, the screening of structural libraries, or any combination of those techniques. Computational software may be used to carry out the sequence alignments and the secondary structure predictions. Structural incoherences, e.g., structural fragments around insertions and deletions, can be modeled by screening a structural library for peptides of the desired length and with a suitable conformation. For prediction of the side chain conformation, a side chain rotamer library may be employed. If the Heφes simplex virus 1 DNA polymerase homolog has been crystallized, the final homology model can be used to solve the crystal structure of the homolog by molecular replacement, as described above. Next, the preliminary model is subjected to energy minimization to yield an energy minimized model. The energy minimized model may contain regions where stereochemistry restraints are violated, in which case such regions are remodeled to obtain a final homology model. The homology model is positioned according to the results of molecular replacement, and subjected to further refinement including molecular dynamics calculations.

RATIONAL DRUG DESIGN

Computational techniques can be used to screen, identify, select and/or design chemical entities capable of associating with Heφes simplex virus 1 DNA polymerase or structurally homologous molecules. Knowledge of the structure coordinates for Heφes simplex virus 1 DNA polymerase permits the design and/or identification of synthetic compounds and/or other molecules which have a shape complementary to the conformation of the Heφes simplex virus 1 DNA polymerase active site. In particular, computational techniques can be used to identify or design chemical entities, such as modifiers, agonists and antagonists, that associate with a Heφes simplex virus 1 DNA polymerase active site or an Heφes simplex virus 1 DNA polymerase-like active site. Potential modifiers may bind to or interfere with all or a portion of an active site(s) of Heφes simplex virus 1 DNA polymerase, and can be competitive, non- competitive, or uncompetitive inhibitors; or interfere with multimeric assemble of polymerase molecules or the interaction of other related proteins with the polymerase. Once identified and screened for biological activity, these inhibitors/agonists/antagonists may be used therapeutically or prophylactically to block Heφes simplex virus 1 DNA polymerase activity and, thus, prevent the onset and/or further progression of Heφes infection. Structure-activity data for analogues of potential modifiers that bind to or interfere with Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active sites can also be obtained computationally. The term "chemical entity," as used herein, refers to chemical compounds, complexes of two or more chemical compounds, and fragments of such compounds or complexes. Chemical entities that are determined to associate with Heφes simplex virus 1 DNA polymerase are potential drug candidates. Data stored in a machine-readable storage medium that displays a graphical three-dimensional representation of the structure of Heφes simplex virus 1 DNA polymerase or a structurally homologous molecule, as identified herein, or portions thereof may thus be advantageously used for drug discovery. The structure coordinates of the chemical entity are used to generate a three- dimensional image that can be computationally fit to the three-dimensional image of Heφes simplex virus 1 DNA polymerase or a structurally homologous molecule. The three-dimensional molecular structure encoded by the data in the data storage medium can then be computationally evaluated for its ability to associate with chemical entities. When the molecular structures encoded by the data is displayed in a graphical three-dimensional representation on a computer screen, the protein structure can also be visually inspected for potential association with chemical entities.

One embodiment of the method of drug design involves evaluating the potential association of a known chemical entity with Heφes simplex virus 1 DNA polymerase or a structurally homologous molecule, particularly with a Heφes simplex virus 1 DNA polymerase active site or Heφes simplex virus 1 DNA polymerase-like active site. The method of drug design thus includes computationally evaluating the potential of a selected chemical entity to associate with any of the molecules or molecular complexes set forth above. This method includes the steps of: (a) employing computational means to perform a fitting operation between the selected chemical entity and an active site or a site nearby the active site of the molecule or molecular complex; and (b) analyzing the results of said fitting operation to quantify the association between the chemical entity and the active site.

In another embodiment, the method of drug design involves computer- assisted design of chemical entities that associate with Heφes simplex virus 1 DNA polymerase, its homologs, or portions thereof. Chemical entities can be designed in a step-wise fashion, one fragment at a time, or may be designed as a whole or "de novo."

To be a viable drug candidate, the chemical entity identified or designed according to the method must be capable of structurally associating with at least part of a Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active sites, and must be able, sterically and energetically, to assume a conformation that allows it to associate with the Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active site. Non-covalent molecular interactions important in this association include hydrogen bonding, van der Waals interactions, hydrophobic interactions, and electrostatic interactions. Conformational considerations include the overall three-dimensional structure and orientation of the chemical entity in relation to the active site, and the spacing between various functional groups of an entity that directly interact with the Heφes simplex virus 1 DNA polymerase-like active site or homologs thereof.

Optionally, the potential binding of a chemical entity to a Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active site is analyzed using computer modeling techniques prior to the actual synthesis and testing of the chemical entity. If these computational experiments suggest insufficient interaction and association between it and the Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active site, testing of the entity is obviated. However, if computer modeling indicates a strong interaction, the molecule may then be synthesized and tested for its ability to bind to or interfere with a Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active site. Binding assays to determine if a compound (e.g., an inhibitor) actually interferes with Heφes simplex virus 1 DNA polymerase can also be performed and are well known in the art. Binding assays may employ kinetic or thermodynamic methodology using a wide variety of techniques including, but not limited to, microcalorimetry, circular dichroism, capillary zone electrophoresis, nuclear magnetic resonance spectroscopy, fluorescence spectroscopy, and combinations thereof.

One method for determining whether a modifier binds to a protein is isothermal denaturation. This method includes taking a sample of a protein (in the presence or absence of substrates) at a fixed elevated temperature where denaturation of the protein occurs in a given time frame, adding the chemical entity to the protein, and monitoring the rate of denaturation. If the chemical entity does bind to the protein, it is expected that the rate of denaturation would be slower in the presence of the chemical entity than in the absence of the chemical entity. For example, this method has been described in Epps et al., Anal. Biochem., 292:40-50 (2001).

One skilled in the art may use one of several methods to screen chemical entities or fragments for their ability to associate with a Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active site. This process may begin by visual inspection of, for example, a Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active site on the computer screen based on the Heφes simplex virus 1 DNA polymerase structure coordinates listed in Table 1 or other coordinates which define a similar shape generated from the machine-readable storage medium. Selected fragments or chemical entities may then be positioned in a variety of orientations, or docked, within the active site. Docking may be accomplished using software such as QUANTA and S YB YL, followed by energy minimization and molecular dynamics with standard molecular mechanics force fields, such as CHARMM and AMBER.

Specialized computer programs may also assist in the process of selecting fragments or chemical entities. Examples include GRID (Goodford, J. Med. Chem., 28:849-57 (1985); available from Oxford University, Oxford, UK); MCSS (Miranker et al., Proteins: Struct. Fund. Gen., 11:29-34 (1991); available from Molecular Simulations, San Diego, CA); AUTODOCK (Goodsell et al., Proteins: Struct. Funct. Genet., 8:195-202 (1990); available from Scripps Research Institute, La Jolla, CA); and DOCK (Kuntz et al., J. Mol. Biol, 161:269-88 (1982); available from University of California, San Francisco, CA). Once suitable chemical entities or fragments have been selected, they can be assembled into a single compound or complex. Assembly may be preceded by visual inspection of the relationship of the fragments to each other on the three-dimensional image displayed on a computer screen in relation to the structure coordinates of Heφes simplex virus 1 DNA polymerase. This would be followed by manual model building using software such as QUANTA or SYBYL (Tripos Associates, St. Louis, MO). Useful programs to aid one of skill in the art in connecting the individual chemical entities or fragments include, without limitation, CAVEAT (P.A. Bartlett et al., in "Molecular Recognition in Chemical and Biological Problems," Special Publ., Royal Chem. Soc, 78:182-96 (1989); Lauri et al., J. Comput. Aided Mol. Des., 8:51-66 (1994); available from the University of California, Berkeley, CA); 3D database systems such as ISIS (available from MDL Information Systems, San Leandro, CA; reviewed in Martin, J. Med. Chem., 35:2145-54 (1992)); and HOOK (Eisen et al., Proteins: Struc., Funct., Genet, 19:199-221 (1994); available from Molecular Simulations, San Diego, CA). Heφes simplex virus 1 DNA polymerase binding compounds maybe designed "de novo" using either an empty active site or optionally including some portion(s) of a known modifier(s). There are many de novo potential modifier design methods including, without limitation, LUDI (Bohm, J. Comp. Aid. Molec. Design., 6:61-78 (1992); available from Molecular Simulations Inc., San Diego, CA); LEGEND (Nishibata et al., Tetrahedron, 47:8985 (1991); available from Molecular Simulations Inc., San Diego, CA); LeapFrog (available from Tripos Associates, St. Louis, MO); and SPROUT (Gillet et al., J. Comput. AidedMol. Design, 7:127-53 (1993); available from the University of Leeds, UK). Once a compound has been designed or selected by the above methods, the efficiency with which that entity may bind to or interfere with a Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase- like active site may be tested and optimized by computational evaluation. For example, an effective Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active site modifier must preferably demonstrate a relatively small difference in energy between its bound and free states (i.e., a small deformation energy of binding). Thus, the most efficient Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active site modifiers should preferably be designed with a deformation energy of binding of at most about 10 kcal/mole; more preferably, at most 7 kcal/mole. Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active site modifiers may interact with the active site in more than one conformation that is similar in overall binding energy. In those cases, the deformation energy of binding is taken to be the difference between the energy of the free entity and the average energy of the conformations observed when the modifier binds to the protein.

An entity designed or selected as binding to or interfering with a Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase- like active site may be further computationally optimized so that in its bound state it would preferably lack repulsive electrostatic interaction with the target enzyme and with the surrounding water molecules. Such non-complementary electrostatic interactions include repulsive charge-charge, dipole-dipole, and charge-dipole interactions.

Specific computer software is available in the art to evaluate compound deformation energy and electrostatic interactions. Examples of programs designed for such uses include: Gaussian 94, revision C (M.J. Frisch, Gaussian, Inc., Pittsburgh, PA (1995)); AMBER, version 4.1 (P.A. Kollman, University of California at San Francisco, (1995)); QUANTA/CHARMM (Molecular Simulations, Inc., San Diego, CA (1995)); Insight U/Discover (Molecular Simulations, Inc., San Diego, CA (1995)); DelPhi (Molecular Simulations, Inc., San Diego, CA (1995)); and AMSOL (Quantum Chemistry Program Exchange, Indiana University). These programs may be implemented, for instance, using a Silicon Graphics workstation such as an Indigo with "IMPACT" graphics. Other hardware systems and software packages will be known to those skilled in the art.

Another approach encompassed by this invention is the computational screening of small molecule databases for chemical entities or compounds that can bind in whole, or in part, to a Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active site. In this screening, the quality of fit of such entities to the active site may be judged either by shape complementarity or by estimated interaction energy (Meng et al., J Comp. Chem., 13:505-24 (1992)).

This invention also enables the development of chemical entities that can isomerize to short-lived reaction intermediates in the chemical reaction of a substrate or other compound that interferes with or with Heφes simplex virus 1 DNA polymerase. Time-dependent analysis of structural changes in Heφes simplex virus 1 DNA polymerase during its interaction with other molecules is carried out. The reaction intermediates of Heφes simplex virus 1 DNA polymerase can also be deduced from the reaction product in co-complex with Heφes simplex virus 1 DNA polymerase. Such information is useful to design improved analogues of known Heφes simplex virus 1 DNA polymerase modifiers or to design novel classes of potential modifiers based on the reaction intermediates of the Heφes simplex virus 1 DNA polymerase and modifier co- complex. This provides a novel route for designing Heφes simplex virus 1 DNA polymerase modifiers with both high specificity and stability.

Yet another approach to rational drug design involves probing the Heφes simplex virus 1 DNA polymerase crystal of the invention with molecules including a variety of different functional groups to determine optimal sites for interaction between candidate Heφes simplex virus 1 DNA polymerase modifiers and the protein. For example, high resolution x-ray diffraction data collected from crystals soaked in or co-crystallized with other molecules allows the determination of where each type of solvent molecule sticks. Molecules that bind tightly to those sites can then be further modified and synthesized and tested for their Heφes simplex virus 1 DNA polymerase modifier activity (Travis, Science, 262:1374 (1993)).

In a related approach, iterative drug design is used to identify modifiers of Heφes simplex virus 1 DNA polymerase. Iterative drug design is a method for optimizing associations between a protein and a compound by determining and evaluating the three-dimensional structures of successive sets of protein/compound complexes. In iterative drug design, crystals of a series of protein/compound complexes are obtained and then the three-dimensional structures of each complex is solved. Such an approach provides insight into the association between the proteins and compounds of each complex. This is accomplished by selecting compounds with inhibitory activity, obtaining crystals of this new protein/compound complex, solving the three-dimensional structure of the complex, and comparing the associations between the new protein/compound complex and previously solved protein/compound complexes. By observing how changes in the compound affected the protein/compound associations, these associations may be optimized. A compound that is identified or designed as a result of any of these methods can be obtained (or synthesized) and tested for its biological activity, e.g., inhibition of Heφes simplex virus 1 DNA polymerase activity.

PHARMACEUTICAL COMPOSITIONS (MODIFIERS) Pharmaceutical compositions of this invention include a potential modifier of Heφes simplex virus 1 DNA polymerase activity identified according to the invention, or a pharmaceutically acceptable salt thereof, and a pharmaceutically acceptable carrier, adjuvant, or vehicle. The term "pharmaceutically acceptable carrier" refers to a carrier(s) that is "acceptable" in the sense of being compatible with the other ingredients of a composition and not deleterious to the recipient thereof. Optionally, the pH of the formulation is adjusted with pharmaceutically acceptable acids, bases, or buffers to enhance the stability of the formulated compound or its delivery form.

Methods of making and using such pharmaceutical compositions are also included in the invention. The pharmaceutical compositions of the invention can be administered orally, parenterally, by inhalation spray, topically, rectally, nasally, buccally, vaginally, or via an implanted reservoir. Oral administration or administration by injection is preferred. The term parenteral as used herein includes subcutaneous, intracutaneous, intravenous, intramuscular, intra- articular, intrasynovial, intrasternal, intrathecal, intralesional, and intracranial injection or infusion techniques.

Dosage levels of about 0.01 to about 100 mg/kg body weight per day, preferably of about 0.5 to about 75 mg/kg body weight per day of the Heφes simplex virus 1 DNA polymerase inhibitory compounds described herein are useful for the prevention and treatment of Heφes simplex virus 1 DNA polymerase mediated disease. Typically, the pharmaceutical compositions of this invention will be administered about 1 to about 5 times per day or alternatively, as a continuous infusion. Such administration can be used as a chronic or acute therapy. The amount of active ingredient that may be combined with the carrier materials to produce a single dosage form will vary depending upon the host treated and the particular mode of administration. A typical preparation will contain about 5% to about 95% active compound (w/w). Preferably, such preparations contain about 20% to about 80% active compound. In order that this invention be more fully understood, the following examples are set forth. These examples are for the puφose of illustration only and are not to be construed as limiting the scope of the invention in any way.

EXAMPLES

EXAMPLE 1

CONSTRUCTION OF FUSION PROTEIN Materials and Methods Reagents. The vector pMAL-CRI, cross-linked amylose, and antibodies to the maltose binding protein were purchased from New England Biolabs (Beverly, MA). Restriction enzymes and other DNA modifying enzymes were purchased from Bethesda Research Labs (Bethesda, MD). Buffers and gels for electrophoresis of proteins were purchased from Integrated Separation Systems (Natick, MA). The [¹⁴C]-labeled protein standards (CFA.756), [methyl-³H]-Thymidine 5 -triphosphate (TRK.424) 40-60 Ci/mmol, and [ S]-methionine (SJ-1015) were purchased from Amersham Biosciences (Piscataway, NJ). The Protein A Sepharose, poly (dA) oligo (dT) template (#27-7868) and the nucleotides were purchased from Pharmacia Coφoration (Peapack, NJ). The protease inhibitors: pepstatin, aprotinin, and leupeptin were purchased from Boehringer-Mannheim (Indianapolis, IN). The amplitaq utilized for polymerase chain reactions (PCR) was purchased from PerkinElmer (Wellesley, MA). The plasmid, pVL 941 was purchased from Invitrogen (Carlsbad, CA). Media required for growth and labeling of Sf9 cells were purchased from BRL-Gibco (Rockville, MD).

Construction of Recombinant Viruses. The E. coli expression vector, pMAL-CRI (Figure 6) was digested with the enzyme Asel, to isolate the sequences encoding the maltose binding protein, multiple cloning site, and part of the lac Z gene. The Asel fragment was treated with Klenow. The baculovirus expression vector pAc373 was linearized with BamHI and treated with Klenow. The Asel fragment from pMAL-CRI was then blunt end ligated into pAc373 to generate the plasmid, pAcMAL (SEQ ID NO:5).

DNA encoding the DNA polymerase from HSV (HSV map. coordinates .413 to .439) was inserted into pAcMAL by digesting the plasmid pDP4 (a gift from D. Coen, Harvard) with Bell and Xbal. The Bell restriction site was located 11 bases upstream from the ATG (Marcy et al., J. of Virol, 64:2208-2216 (1990)). The fragment was inserted into the BamHI-Xbal site of pAcMAL This plasmid was designated as DP2.

Sf9 cells were cotransfected using calcium phosphate precipitation with wild type AcNPV DNA and transfer plasmids containing either the MBP, the DNA polymerase gene from HSV-1, or the MBP-fusion proteins (Summer et al., ".4 Manual of Methods for Baculovirus Vectors and Insect Cell Culture Procedures, " College Station: Texas Agricultural Experiment Station (1987)). The recombinants were isolated by selecting "inclusion negative" plaques. The virus was plaque purified a total of 4 times. Construct of the modified linker between MBP and HSV 1 POL. The region surrounding the desired mutation as shown in Figure 8 (nucleotides 4607- 6080) was excised from pACMAL-HSV 1 POL vector with BgllJ and ligated into the vector pQE60. Mutagenesis was performed using desired mutagenic oligonucleotides in the Stratagene Quickchange mutagenesis system (La Jolla, CA). DNA sequence analysis confirmed that the correct mutations have been made. The Bgiπ fragment was excised from pQE60 and ligated back into the Bgiπ site of the pACMAL-HSV 1 POL. The BgllJ fragments were confirmed to be in the correct orientation by restriction mapping, and DNA sequence analysis onfirmed that the constructs were correct.

Cell Localization. Sf9 cells (2 x 10 cells) were infected with either baculovirus containing BP58 (HSV DNA polymerase without the maltose binding protein, obtained from D. Coen), DP2, or MAL. The cells were labeled 48 hours post-infection in Graces methionine free medium containing 100 microCi/ml [³⁵S]-methionine for 2 hours and harvested by pelleting the cells. The cells were resuspended in 1 ml of cold buffer A (20 mM Hepes pH 7.6, 0.5 mM DTT, 10 MM NaHS0₃, and 0.5 mM PMSF) (Marcy et al., Nucl. Acid Res., 18:1207-1215 (1990)) and broken in a dounce homogenizer. The nuclei and cellular debris were removed by centrifugation for 10 minutes at 3000 φm in a Sorvall RT 6000B centrifuge. The cytosolic fraction (fraction II) was removed for analysis. The nuclei and debris were resuspended in 500 microliters of buffer A and an additional 500 microliters of buffer A containing 3.4 M NaCI was added. The insoluble proteins (fraction III) were removed by a high speed spin in a Tomy centrifuge. The soluble nuclear material (fraction U) was removed for analysis. The fraction HI material was solubilized using 1 ml of RIPA buffer (20 mM Tris pH 8.0, 100 mM NaCI, 1 mM EDTA, 1.0% deoxycholate, 1.0% NP40, and 0.1% SDS) and mixed for 5 minutes. Antibodies (25 microliters) to the truncated form of HSV DNA polymerase were added to each sample and allowed to bind overnight at 4°C The samples were incubated with 50 microliters of a 115 micrograms/ml suspension of Protein A Sepharose for 2 hours at room temperature. The samples were centrifuged and the pellets washed repeatedly with RIPA buffer. The protein was boiled in protein sample buffer containing SDS and BME, and electrophoresed on a SDS-PAGE. The gel was fixed and treated with Enhance (NEN-Dupont).

Results

Construction of Recombinant Baculovirus Vectors. The pAcMAL vector (SEQ ID NO:5) was constructed by inserting the Asel restriction fragment from pMAL-CRI into the BamHI site of pAc373. The Asel fragment contained the malE, Lac Z, and multiple cloning site of pMAL-CRI vector (Figure 6). The polyhedrin promoter from pAc373 was used in the baculovirus system for expression of the M13P fusions. The HSV DNA polymerase was cloned in-frame into the multiple cloning site of pAcMAL at the BamHI Xbal site to generate pAcMAL-DP2. The MBP-fusion should add about 40 kDa to the molecular weight of the protein resulting in a predicted molecular weight of 175 kDa for the fusion protein.

The pAcMAL and pAcMAL-DP2 vectors were used to isolate recombinant baculoviruses expressing MBP and MBP-HSV polymerase, respectively, as described in Materials and Methods. Cell Localization. Immunoprecipitation experiments using

[³⁵S]methionine labeled Sf9 cells infected with recombinant baculoviruses were used to detect the size and cellular localization of the fusion proteins. Sf9 cells were infected with recombinant baculoviruses and 48 hours post infection the media was removed and replaced with [ Sjmethionine containing media. Two hours later the cells were harvested. The cells were broken by douncing in a low salt buffer (Marcy et al., Nucl. Acid Res., 18:1207-1215 (1990)), and the disrupted cell material was separated into a cytosolic fraction (fraction I), a nuclear fraction (fraction 13), and an insoluble protein fraction (fraction HI) as described in Materials and Methods. A rabbit polyclonal antisera raised against a truncated version of the MBP-HSV DNA polymerase expressed in bacteria (see Materials and Methods) was used to immunoprecipitate HSV POL expressed from BP58 (expresses nonfusion form of HSV POL), DP2 and MAL (expresses only MBP). The rabbit antisera specifically immunoprecipitated a 140 kDa protein from BP58 infected cells, a 175 kDa protein from DP2 infected cells and a 50 kDa protein from MAL infected cells. The gel mobilities of the HSV POL, MBP-HSV POL, and MBP are in agreement to the sizes predicted for these proteins based on there DNA sequence. There are several other bands visible in the immunoprecipitation reactions. These bands may either be contaminants or they may be cleavage products of the HSV DNA polymerase. The HSV DNA polymerase appears in the cytosolic fraction in both fusion (DP2) and nonfusion (BP58) forms. The HSV DNA polymerase fusion protein is made at higher levels than the non-fusion form in baculovirus. The MBP appeared to be cytosolic in baculovirus infected Sf9 cells.

EXPRESSION AND PURIFICATION Materials and Methods

Materials. PI 1 phosphocellulose resin was purchased from Whatman (Clifton, NJ). Bradford protein assay solution and ceramic type II hydroxyapatite (CHT HA) resin were obtained from Bio-Rad (Hercules, CA). HiTrap heparin column, ³H-dTTP (TRK576, 111 Ci/mmol), and streptavidin SPA beads (RFN006) were purchased from Amersham Biosciences (Piscataway, NJ). EDTA-free protease inhibitor cocktail tablets are available under the trade designation COMPLETE (#1873580) from Roche (Basel, Switzerland). ssDNA template and primer strands were from Oligos, Etc (Wilsonville, OR). All other chemicals were purchased from Sigma (St. Louis, MO) with the highest purity.

Protein expression. The MBP-HSV 1 POL was expressed in a baculovirus/insect cell system. To identify the best expression conditions, cell pellets from 500 ml SF9 culture grown 65 hours post infection at different virus multiplicities of infection (MOI) were tested for yields by purification. Large- scale protein expression was realized in a bioreactor with 20-40 liters of baculovirus SF21 or SF9 cell cultures (Kemp Biotech Inc, Frederick, MD). Cells were pelleted by slow speed centrifugation. Protein extraction. A flow-chart of the purification scheme is shown in

Figure 9. All purification steps were performed at 4°C or on ice unless noted. About 10 grams of cell pellets were suspended in 50ml of lysis buffer (lOmM Tris-HCl, pH=7.5, lOmM KCl, 1.5mM MgCl₂), and 1 tablet of COMPLETE protease inhibitor cocktail, 100 microliters lOmg/ml PMSF stock, and 50 microliters IM DTT stock were added. The suspended cells were incubated on ice for 20 minutes and then disrupted by 20 strokes of Dounce homogenization (Wheaton #357546). Solid NaCI was added to the lysates to the final concentration of 1.5M and incubated on a mixer available under the trade designation Hoefer Red Rotor for an hour. The homogenate was centrifuged at 16,500 rpm in a SA600 rotor for an hour to clear cell debris and nuclei. The clarified supernatant was dialyzed twice overnight against each of 2 liters of HEPES buffer (20 mM HEPES, pH=7.5, 10% glycelrol, 1 mM DTT, 0.1 mM EDTA), plus 100 mM NaCI. Particulate precipitated during dialysis was removed by 20 minutes centrifugation at 16,500 rpm followed by filtration through 0.22 micrometer membrane (Corning #430320). To promote the fusion protein's digestion by endogenous proteases, dialysis was done at room temperature and no protease inhibitors were added during cell disruption.

Pll Phosphocellulose column. The following chromatography steps were done either on an Akta or a FPLC chromatography system (Amersham Pharmacia Biotech), with a flow rate of 1 ml/minutes all the time. A phosphocellulose column was packed into an XK26/20 column (Amersham Pharmacia Biotech, with 1 adapter) by treating 4.0 g PI 1 phosphocellulose resin according to manufacture's instructions, resulting a bed volume of 40ml. The column was washed with 200 ml of HEPES buffer + 0.1M NaCI. Filtered dialysate from salt extraction was loaded onto the column. After loading, the column was washed with 180 ml HEPES buffer + 0.1M NaCI. A 250ml gradient from 0.1M to 0.6M NaCI in HEPES buffer was applied to elute the bound protein. Fractions containing MBP-HSV 1 POL identified by SDS-PAGE gel were pooled. Protease digestion of the fusion protein. Digestion of MBP-HSV IPOL by unknown endogenous protease(s) can be done either on crude homogenate before PI 1 column or as a salvage step after the HA column (Figure 9). Before PI 1 column, the digestion occurs during dialysis at room temperature. However, the digestion was usually incomplete, and sometimes the pooled PI 1 fractions needed to be left for several days to achieve better digestion before the HA column. Factor Xa digestion of fusion protein containing two factor Xa sites in the linker was carried out after the PI 1 column step. Pooled PI 1 fractions were combined and 300 μg factor Xa was added and the reaction was carried out at 4°C for 5 hours. The proteolysis was quenched with the addition of phenylmethylsulfonyl fluoride to a final concentration of lmM.

Hydroxyapatite (HA) column. This column proves very useful for separating cleaved and uncleaved fusion protein. After adding 8M guanidine-HCl stock to a final concentration of lOOmM, the digested PI 1 column fraction pool was loaded directly on to a 15ml HA column packed in an XK16/20 column (Amersham Pharmacia Biotech, 1 adapter), equilibrated with low phosphate buffer (lOmM KH₂PO , pH=7.0, lOOmM NaCI, 10% glycerol, 1 mM DTT, lOOmM guanidine-HCl) at the rate of 1 ml/minute. After loading, the column was washed with 50 ml of low phosphate buffer with IM NaCI, followed by 50 ml normal low phosphate buffer to bring back conductivity. Protein was eluted with an 110 ml gradient of 10 mM to 75 mM phosphate (pH=7.0, with other components the same as the low phosphate buffer). Fractions containing truncated HSV 1 POL and full-length fusion protein identified by SDS-PAGE were pooled separately. The column was cleaned with 100 ml 500 mM NaH₂PO4, pH=7.0 buffer at room temperature and re-equilibrated with 100 ml of low phosphate buffer. Heparin column. Pooled HA column fractions containing either the truncated HSV 1 POL or undigested fusion protein were separately loaded onto a 5ml HiTrap heparin column equilibrated with HEPES buffer containing 100 mM NaCI and 100 mM guanidine-HCl. After loading, the column was washed with 100 ml of the same buffer. Proteins were eluted with a 100 ml gradient of 100 mM NaCI to 600 mM NaCI in HEPES buffer with 100 mM guanidine-HCl. Fractions containing HSV 1 POL identified by SDS-PAGE were pooled. The column was cleaned with 1.6M NaCI in HEPES buffer then re-equilibrated with five column volume of starting buffer. Activity assay. HSV 1 POL activity was measured using scintillation proximity assay which measures the incoφoration of tritiated deoxythymidine triphosphate (³H-dTTP) into a DNA poly (dA)₂₅₀- biotinylated-p(dT)₁₆ template/primer duplex. Fractions were diluted at 1:100 ratio into respective wells of a 96-well assay plate in reaction buffer of 7.6 mM HEPES pH 7.5, 5.6 mM MgCl₂ , 45 micrograms/mL BSA, 5 mM CHAPS, 5% DMSO, and 1 mM DTT. The incoφoration reaction was performed at 26.3°C and was initiated by the addition of 100 nM ³H-dTTP and 6 nM template/primer to each well. Incoφoration was allowed to proceed until stopped by the addition of 0.5 M EDTA and streptavidin-conjugated beads containing a scintillant for detection. Captured radioactivity of incoφorated ³H-dTTP was counted in a Packard

Topcount scintillation counter. The activity assay was also performed in parallel with positive and negative controls.

Amino acid analysis. Amino acid analysis of purified HSV 1 POL was carried out. The N-terminal sequence was determined using an automated Edman degradation method. The amino acid composition was carried out using an HC1 hydrolysis method. With this method, tryptophan and cysteine were not determined. Mass spectroscopy experiments on purified protein was also performed. Results

Protein expression. Cell pellets from 500 ml SF9 cell cultures after 65 hours of post-infection growth, with virus MOIs of 0.25, 0.75, 1.5, and 3.0, respectively, were tested for expression level. Cells were disrupted and purified through the PI 1 column. The integrated peak area corresponding to MBP-HSV 1 POL (or truncated HSV 1 POL) was used as quantitative index for the amount of protein expressed. Cell cultures with MOI=0.75 gave the best result compared to samples of other MOIs. The expression level of HSV 1 POL is about 1.5 and 2 times higher than that of MOI=l .5 and 3.0 respectively. The expression level of the MOI=0.25 culture was barely detectable. Large-scale expression (40 liters) of MBP-HSV 1 POL in SF21 cell with a MOI of 0.75 in a bioreactor confirmed significant expression. 15-30 mg purified HSV 1 POL was obtained from cell pellets from 2 liters of culture. PI I column. The first and the major protein peak is eluted between 150 mM and 350 mM NaCI on the PI 1 column. The SDS-PAGE gel of these fractions showed two major protein bands at approximately 170 kilodaltons (KD) and approximately 120 KD positions. These two protein bands exist in the crude lysate but disappear in the PI 1 column flow-through. The 170 KD band is close to the position of the full-length MBP-HSV 1 POL fusion protein and the 120KD band is close to that of the HSV 1 POL (calculated molecular weight 178,688 and 132,654 daltons, respectively). Characterizations outlined below confirmed that the lower band is that of a truncated HSV 1 POL and the upper band is that of the MBP-HSV 1 POL fusion protein. The relative amount of these two proteins depends on cell growth and disruption conditions and varies between batches. With protease inhibitors in the lysis buffer and dialysis at 4°C, almost all MBP-HSV 1 POL is intact. On the other hand, without protease inhibitors and dialysis at room temperature the amount of the truncated HSV 1 POL increases substantially. The fact that these two proteins almost co-elute makes it difficult to separate them on Pi 1 column. DNA polymerase activity assay showed that PI 1 column effectively captures almost all POL activity (data not shown), and fractions containing 170KD and 120KD proteins are highly active. Besides these fractions, proteins eluted later also contain POL activity. Most of the proteins in the later fractions are of much lower molecular weights. Activity in those fractions presumably come from the endogenous insect cell DNA POL, baculovirus DNA POL or smaller fragments of HSV 1 POL. It is preferable to pack a fresh PI 1 column each time to capture the HSV 1 POL from the crude lysate rather than regenerating the column material both because the phosphocellulose resin is cheap and it gets compressed after each run. Protein digestion. Observation of cleavage of the HSV 1 POL during cell disruption and protein purification led to a digestion method relying on the endogenous protease(s). The digestion site was shown to occur N-terminally to N43 of the HSV 1 POL by N-terminal sequence analysis (Figure 8). Amino acid composition and mass spectrum analysis on purified protein showed that the protein is intact from N43 to the C-terminus. Completion of digestion using this method depends on cell disruption and protein extraction conditions. Less than one third of the fusion protein generally gets digested this way before the HA column, even after being left for several days. Digestion by factor Xa of the fusion protein with two factor Xa site in the linker is controlled and the digestion is complete in 5 hours. The N-terminal of the digestion product was confirmed to be N43..

HA column. Because of its high selectivity, the HA column is an effective step in getting rid of most impurities and the only step in separating the proteolytically processed HSV 1 POL from the full-length fusion protein. Contrary to reports on HSV 1 POL's behavior on this resin (Knopf et al.,

"Heφes simplex virus type 1 DNA polymerase: Eukaryotic model enzyme and principal target of antiviral therapy" in Frontiers of Virology, Becker and Darai, eds., 3:87-135 (1994)), the protein is retained at a phosphate concentration of 10 mM. Regardless of what method was used to digest the fusion protein, truncated HSV 1 POL elutes at almost the same phosphate concentration (around 20 mM phosphate). Full-length fusion protein elutes at phosphate concentrations higher than 40 mM. We also took advantage of the special elution profile of the HA column to wash out a lot of impurities using IM NaCI + 10 mM phosphate before the phosphate gradient. This step dramatically improved the purity of the protein. We also found that the specific activity of the truncated protein is much higher than the full-length fusion protein. We also noticed that in the presence of 100 mM guanidine-HCl, HSV 1 POL elutes much more shaφly and symmetrically. Since affinity of a protein toward the HA column depends on its three dimensional structure, we can hypothesize that HSV 1 POL is more homogenous and compact in the presence of 100 mM guanidine-HCl.

HiTrap Heparin column. The heparin column removes some minor impurities in the pooled fractions of the truncated HSV 1 POL from the HA column and at the same time concentrates the protein. The high affinity of HSV 1 POL toward heparin resin ensures that the pooled HA column fractions can be loaded directly without dialysis onto the Heparin column. The presence of 100 mM guanidine-HCl also changes heparin column elution profile. HSV 1 POL elutes at much higher salt concentration in the presence of guanidine than without. This result also suggests that in the presence of 100 mM guanidine-HCl, HSV 1 POL may have a more compact three-dimensional structure thus a better formed DNA binding cleft. Heparin resin is a pseudo-affinity resin toward DNA binding proteins because its poly-sulfated-sugar groups mimic the negatively charged DNAThe amount of the highly purified HSV 1 POL after this column is about 8-10 mg.

Results of amino acid analysis. N-terminal sequence analysis showed that each batch of purified HSV 1 POL has homogeneous N-termini, and is consistent with the digestion method used. The amount of protein with heterogeneous N-terminal sequence is less than 10%. Amino acid composition analysis on N43-C HSV 1 POL showed that it is intact to the C-terminus of the protein, which covers all known function domains, including the 3'-5' proofreading exo-nuclease domain, the 5-3' exo-nuclease domain, the DNA polymerase domain, and the accessory protein binding domain. Mass spectroscopy on N43-C HSV 1 POL (not shown) showed a peak of 132,468 Da, which is within experimental error of the calculated molecular weight of 132,454.

CRYSTALLIZATION PROCEDURE

Protein was prepared by adding 40 μL of HSV POL (25 mg/mL) to 5 μL of DMSO in a warm tube. After rapid mixing, the protein was placed on ice for more than 1 hour. A crystallization screen was set up in a 24 well tray with 1 mL of the following solution in each well: 0.1 M HEPES, pH 7.0; 5 mM DTT; 0.1 M guanidine-HCl; 30-60 mM ammonium sulfate (in 10 mM increments); and 3- 8 % PEG-3350 (in 1 % increments). 1.25 μL of each well solution was added to 1.25 μL of protein and placed on a cover slip over that well according to the hanging-drop method. After 8 days, several wells contained crystals with one crystal edge longer than 0.2 mm.

One of these drops was well C2 (0.1 M HEPES, pH 7.0; 5 mM DTT; 0.1 M guanidine-HCl; 50 mM ammonium sulfate; and 4% PEG-3350). The final cryoprotectant solution used for this drop contained 0.1 M HEPES, pH 7.0; 5 mM DTT; 0.1 M guanidine-HCl; 60 mM ammonium sulfate; 10% PEG-3350; 11% DMSO; and 10% PEG-1000. At 5-minute intervals, the coverslip was inverted and increasing amounts of cryoprotectant were added. The amounts of cryoprotectant were: add 0.25 μL, add 0.25 μL, add 0.5 μL, add 0.5 μL, add 1 μL, add 1 μL, remove 2 μLand add 2 μL, remove 4 μLand add 4 μL, and remove 8 μLand add 8 μL. After these 9 additions of cryoprotectant, the drop was allowed to sit for at least 30 minutes. Crystals were looped out and plunged into liquid nitrogen and stored for data collection.

Equivalent crystals grow for a variety of constructs. Crystallization of single Xa and dual Xa fusion protein is expected using the same procedure to give similar crystals. X-RAY DIFFRACTION CHARACTERIZATION

Data Collection: All data collection was carried out at the APS synchrotron at beamline EVICA 17DD using synchrotron radiation. The crystals were kept at temperature of 100K under constant flow of cold N2 stream. To collect complete data set, 120 frames of 1° oscillation images were collected for each data set. For multiple wavelength experiments, energy scans were done on frozen crystals and relevant absoφtion edges were selected for particular heavy atoms. For multiple wavelength experiments, both direct and inverse beam for each wavelength were collected, and to reduce systematic errors due to radiation damage, data collection was done in interleaved 30° segments. X-ray diffraction images were recorded on an ADSC-Quantum210 CCD detector, and were processed with program HKL2000 (Otwinowski, et al., in Macromolecular Crystallography, Part A, Carter & Sweet eds, Academic Press, pp. 307-326 (1997)). Different data sets were scaled together with programs from CCP4 package (Collaborative Computing Project #4, Acta Cryst., D50:760-763 (1994).

Structure solution: The structure was solved by multiple anomalous dispersion method using crystals formed by selenomethionine incoφorated protein and soaked with 20mM trimethyllead acetate overnight. 37 out of possible 40 selenomethionine sites and 12 lead sites were located with program SHELXD (Sheldrick et al., Acta Cryst. B5L423-431 (1995)). Heavy atom refinement and phasing were carried out with program MLPHARE (Otwinowski, in Isomorphous Replacement and anomalous Scattering, Wolf, Evans & Leslie eds (Warrington, UK, Daresbury Laboratory), P80-86 (1991)). Initial model was built into the 3.5 A experimental phased electron density map with program O (Jones et al., Acta Cryst. A47: 110-119 (1991)). Location of fragments of other polymerases at the experimental map with program MOLREP (Vagin et al., J. Appl. Cryst., 40:1022-1025 (1997)) helped registration of amino acid residues. The structure refinement were carried out with program CNX (Brunger, Methods Mol. Biol., 56:245-266 (1996)) or REFMAC5 (Murshudov et al., Acta Crystl., D53:240-255 (1997)), and manually rebuilt with program O, against 2.8A resolution data collected on a native crystal. The solvent content of the crystal was 55%. The Matthews coefficient was 2.8 A/Da.

TABLE 8: Data collection, phasing and refinement Statistics

-

Native SeMet wavelength 1 SeMet wavelength 2

(data set ID aps 1029) (data set-D aps 1337) (data setID apsl337)

Space Group P2ι2ι2ι P2,2ι2, P2ι2j2ι

Unit cell constants 103.92, 125.55, 220.58 105.11, 123.96, 217.67 105.11, 123.96, 220.58, 217.67

Wavelength 1.0 0.9794 0.9504

Resolution 31.4-2.8 54-3.2 54-3.2

Observed reflections 781,296 897,759 446,446

Independent reflections 81,297 47,478 46,949

I/σ(D 30.0(2.4) 7.8(2.2) 7.4(1.8)

Average redundancy 9.4 (8.2) 18.9(19.1) 9.4(9.5)

Rmerge 0.071 (0.64) 0.078 (0.331) 0.086(0.411)

Completeness % 100 (100) 99.2(99.9) 99.1(99.9)

Rcullis (to 3.5 A) - 0.87 (0.82) -

Rcullis, anom (to 3.5 A) - 0.70 0.77

Rfactor 0.245 (0.264) - -

Rfree 0.331 (0.351) - -

The complete disclosure of all patents, patent applications including provisional applications, and publications, and electronically available material (e.g., GenBank amino acid and nucleotide sequence submissions; and protein data bank (pdb) submissions) cited herein are incoφorated by reference. The foregoing detailed description and examples have been given for clarity of understanding only. No unnecessary limitations are to be understood therefrom. The invention is not limited to the exact details shown and described; many variations will be apparent to one skilled in the art and are intended to be included within the invention defined by the claims.

SEQUENCE LISTING FREE TEXT

SEQ ID NO: 1 Amino acid sequence of HSV-1 strain KOS DNA polymerase

SEQ ID NO:2 Nucleotide sequence for a gene that encodes HSV-1 strain KOS DNA polymerase

SEQ ID NO:3 Amino acid sequence of Maltose-HSV-1 strain KOS DNA polymerase fusion protein

SEQ ID NO:4 Nucleotide sequence for a gene that encodes Maltose- HSV-1 strain KOS DNA polymerase fusion protein

SEQ ID NO:5 Nucleotide sequence for the pACMAL vector.

TABLE 1: Structure Coordinates

ATOM 1 N GLY A 60 76 .784 75. .051 12. .411 1. .00 71, .85 N

ATOM 3 CA GLY A 60 76 .202 75. .172 11. .026 1. .00 71, .77 C

ATOM 6 C GLY A 60 75 .469 73. .939 10. .476 1. .00 71, .18 C

ATOM 7 0 GLY A 60 75. .918 73. .355 9. .489 1. .00 71, .30 O

ATOM 10 N PRO A 61 74, .366 73. .504 11. .103 1. .00 69, .80 N

ATOM 11 CA PRO A 61 73, .607 72. .377 10. .550 1. .00 68, .65 C

ATOM 13 CB PRO A 61 72. .410 72. .265 11. .494 1. .00 68. .33 C

ATOM 16 CG PRO A 61 72, .816 72, .906 12. .725 1. .00 68. .92 C

ATOM 19 CD PRO A 61 73, .793 73. .983 12. .377 1. .00 69. .27 C ATOM 22 C PRO A 61 73, .164 72. .594 9. .057 1. .00 68. .33 C

ATOM 23 0 PRO A 61 72, .714 73. .704 8. .643 1. .00 67. .42 O

ATOM 24 N THR A 62 73, .319 71. .522 8. .257 1. ,00 67. .52 N

ATOM 26 CA THR A 62 72. .839 71. .509 6. .856 1. .00 66. .81 C

ATOM 28 CB THR A 62 73. .427 70. .216 6. .036 1. .00 66. .84 C

ATOM 30 OG1 THR A 62 73. .467 69. .037 6. .850 1. .00 63. .43 O

ATOM 32 CG2 THR A 62 74. .988 70. .449 5. .582 1. .00 65. .64 C

ATOM 36 C THR A 62 71, .269 71. .757 6. .771 1. .00 66. .35 C

ATOM 37 0 THR A 62 70, .784 72. .557 5. ,937 1. .00 66 . .84 O

ATOM 38 N GLN A 63 70, .514 71. .112 7. .682 1. .00 65. ,01 N

ATOM 40 CA GLN A 63 69, .092 71. .422 7. .959 1. ,00 63. .35 C

ATOM 42 CB GLN A 63 68. .219 70. .159 7. .923 1. ,00 63. ,00 C

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CM CM CM CM CM H CM CM CM CM CM CM H CM CM CM CM CM CM CM CM CM I 1 1 1 I 1 1 1 1 1 I 1 1 1 1 1 1 1 1 1 1 cn o VD I^s- VD cn co ro ro ro cn rH CM m H cn H m 00 ro o CM o VD ro I^s- 00 ro VD ro CM H ro m sμ CO ro H sμ ro in I^s- oo ro en I^s- o O CM sμ CM CM ro in ro ro sμ I- cn r- co VD sμ CM sμ cn

H H CM CM CM H ro sμ ro sμ sμ m sμ sμ in ro ro CM CM CM CM ro

LO m m in in in in in in m m m in m m m in m in m in in

VD on I^s- vo CM in CM sμ I^s- ro m H cn on cn o ro VD in vo oo CM vo CM ro I^s- ro m sμ CM r- sμ H m cn cn H H r- in VD oo I^s- H sμ CO vo I^s- in ro H m ΓO CM sμ in co I^s- o o CM o ro ro sμ cn sμ ro CM r-i o CM in sμ VD I- oo cn I- I^s- co ∞ 00 on o H o o sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ Lf) in in sμ

H H v-i rH H H v-i H CM CM CM CM CM CM CM ro ro ro ro ro ro ro

H H H H H H v-i rH H H H rH H H H H v- H H H H vi

CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM ft fiϋ fit fit fiϋ fi fit fit fiϋ ft fit ft ft fit fit fit fiϋ fit fiϋ fit fit fit

P P P P P P P P μQ μQ μQ μQ μQ μQ μQ ft ft ft ft ft ft ft μQ μQ μQ μQ μQ μQ μQ μQ fit, fiϋ fiϋ fit fit fit, to to to co to to CO

0 0 0 0 0 0 0 0 > > > > > > > fiϋ fit fiϋ fit fiϋ fiϋ fit

H CM H CM H CM ft PQ CD ou PQ PQ fit, PQ CD CD fit PQ CD P P

U U U O O U O a u U u U U o a u U U o o U

H ro vo cn σ H CM ro sμ vo oo o sμ co en o CM sμ t- co cn o

I^s- Γ- I^s- I^s- oo oo oo oo 00 oo oo en en on cn o O o o o o H ro ΓO ro ro ro ro ro ro ro ro ro ro ro CO ro sμ sμ sμ sμ sμ sμ sμ

CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 O O O O O O O O O O O O O O O O O O O O O O E^H H EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH fi fi fi fi fi fiϋ fi fi fiϋ fi fiϋ fiϋ fi fi fi fi fiϋ fi fi fi fiϋ fiϋ ATOM 2411 0 ASP A 213 46.430 53.006 -27.294 1.00111.35 O

ATOM 2412 N ARG A 214 46.266 55.062 -26.445 1.00114.16 N

ATOM 2414 CA ARG A 214 45.062 55.452 -27.190 1.00116.61 C

ATOM 2416 CB ARG A 214 43.795 54.828 -26.590 1.00117.06 C

ATOM 2419 CG ARG A 214 43.101 53.796 -27.450 1.00117.62 C

ATOM 2422 CD ARG A 214 41.701 53.528 -26.972 1.00119.16 C

ATOM 2425 NE ARG A 214 41.243 52.185 -27.310 1.00120.77 N

ATOM 2427 CZ ARG A 214 39.991 51.752 -27.147 1.00120.70 C

ATOM 2428 NH1 ARG A 214 39.062 52.562 -26.646 1.00119.73 N

ATOM 2431 NH2 ARG A 214 39.670 50.500 -27.490 1.00120.93 N ATOM 2434 C ARG A 214 44.944 56.977 -27.136 1.00118.42 C

ATOM 2435 0 ARG A 214 44.436 57.604 -28.086 1.00118.97 O

ATOM 2436 N HIS A 215 45.377 57.541 -25.995 1.00120.18 N

ATOM 2438 CA HIS A 215 45.617 58.984 -25.804 1.00121.37 C

ATOM 2440 CB HIS A 215 45.215 59.422 -24.374 1.00121.71 C

ATOM 2443 CG HIS A 215 44.219 58.503 -23.722 1.00124.16 C

ATOM 2444 ND1 HIS A 215 42.864 58.558 -23.987 1.00127.04 N

ATOM 2446 CE1 HIS A 215 42.236 57.623 -23.293 1.00127.56 C

ATOM 2448 NE2 HIS A 215 43.137 56.948 -22.602 1.00127.39 N

ATOM 2450 CD2 HIS A 215 44.385 57.474 -22.856 1.00126.04 C

ATOM 2452 C HIS A 215 47.111 59.202 -26.089 1.00121.61 C

ATOM 2453 0 HIS A 215 47.939 59.154 -25.177 1.00121.54 O

a u u u u u u o a u u u u o a u o a u u co u

en sμ o in oo CM I^s- H cn o sμ CM 00 m cn ro H vo I^s- oo sμ vo o ro o D sμ r-i rH on o vo VD ro CM CM o CM on o ro o sμ

CM CM CM CM CM CM CM H CM H H H o oo o r-i H on on on o I^s- CM CM CM CM CM CM CM CM CM CM CM CM CM rH CM CM CM H H H CM H H H H rH r- H H v- H H rH H H rH H rH rH H H rH H rH O O O O o O o o O o O o o O o O O o o O O O O O CD o o o o o o o O o o O o O O o o o O O

H H H H H H rH rH H H H rH H H H H H H rH H rH H m I^s- ro H cn ro o CM en sμ 00 ro CM irirVO cn VD O VD sμ in

I^s- I^s- H ro in H CM I^s- vo H I^s- oo CM on on I^s- H vo CM VD ro co ro en sμ vo I^s- H cn oo o CM ιn on sμ H oo O sμ vo VD VD ro rH r- r- r- I^s- VD cn I- VD on en o o H CM o cn 00 on on O 00

CM CM CM CM CM CM CM CM CM CM ro ro co co ro CM CM CM CM ro ro CM 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 co ro o vo I- H cn sμ cn vo I^s- co o ro CM ro o o sμ oo rH O

H H t- oo vo ro sμ vo in ro o sμ 00 CM r- H r-i CM I^s- vo vo on sμ ro ro oo I^s- ro oo ro v-i I^s- H CM VD ro rH I^s- VD I^s- VD vo ro on on cn o rH CM CM I^s- I^s- r- m m in VD r- ! m vo sμ sμ in m sμ in in VD VO VO vo m m m m m in m m m in in in m in m in cn vo ro on m v-i in CM CM I^s- sμ o I^s- m o in CM CM I^s- t- ro o rH I- r- r- on vo H in sμ sμ on cn co ∞ r- t- sμ m CM CM H H sμ r- I^s- sμ ro m ro I^s- CM VD H VD CM on H rH I- oo ro cn vo co

I^s- co cn on o on on cn cn on on I^s- t- vo H H H ro ro in ro sμ sμ sμ sμ in sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ in m in m in in in

vo o vo vo VD vo VO o I^s- I^s- I- I^s- I^s- I^s- r- I- t- CO co co oo oo

H H H H H H rH H H H H v-i H v-i H H H H H H H rH

CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM fit fit fiϋ fit fiϋ fiϋ fit fiϋ fit fiϋ fit fit fit fit f fiϋ fit ft fiϋ ft fit ft

P P P P P P P P

PQ PQ PQ PQ PQ PQ PQ aμQ aμQ aμ a a a a a a to to co CO co

PQ Q μQ μQ Q μQ μQ μQ H H H tx H μQ μQ μQ μQ μQ μQ μQ μQ 0 0 0 0 0 0 0 0 0 u u U u u CM H CM a fiϋ PQ CD P P fiϋ PQ CD P PQ PQ fiϋ PQ CD

U U u u u u o a u u u u O a U O a u U to u sμ vo 00 H ro I^s- H CM ro m I^s- o ro sμ L0 oo cn o CM sμ I^s- 00 in m in vo VD VO I^s- r- I^s- I^s- I- oo oo oo oo oo oo cn cn cn cn cn sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ

CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 O O O O O O O O O O O O O O O O O O O O O O EH EH EH EH EH EH H EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH ft fit fit fit fit fit fit fit fit fit fit fit fit fit fit ft fit fit fit fit fit fit o a u u u u a u a a u o a u u u o a u u u u

I- H sμ CM H sμ in vo ro oo o o v- I^s- cn o ro on vo I^s- .ro in ro ro t- H VD sμ CM CM CM I- r-i sμ I^s- CM vo co CD vo o o o I^s- r- in ro sμ in r- cn en en ∞ CM CM o o o on cn on on cn cn cn

H vi H H H t-H H H H v-i H rH H H rH o o o o o o o

H rH v-i H H H H rH H H H rH rH -i rH rH v-i H H rH H rH

O O o O o o o o o O O O O O O O o o o O o O

O o o O o o o o o O O O O O O O o o o O o O v-i H H H H H H H H H H r-i H v-i H H H H H H r-i H vo 00 CM VO CM in sμ m ro m sμ sμ VD sμ en VO CM ro ro O cn VO iron m CM iroo o I^s- σn o vo I^s- 00 ro H m I^s- sμ VD ro ro on on H oo O on v-i ro VD co oo ro ro O I- cn CM O H I^s- oo o m r- r- m m sμ sμ sμ ro CM ro m sμ vo in o m sμ VO m VD co IT-

CM

O CM o ∞ I^s- ro 00 vo H vo m sμ I^s- H CM sμ o I^s- vo I^s- H o r- in H in I- I^s- ro sμ sμ VD o vo 00 CO cn CM s sμ ro O m ro m vo cn ro co sμ cn irCM O VD VD in ro ro oo VD sμ o H in vo in sμ sμ m vo I- oo on cn v-i ro ro CM H o H H CM ro sμ ro CM in m in in ω m in m in VD m in m in in m m in m m m m co vo CM v-i oo H o H en sμ CM sμ m sμ CM oo cn CM o sμ r- vo r- on I^s- on rH sμ r-i m o H m CM m sμ CM r-i cn sμ ro sμ m sμ co cn ro rH H cn cn o H H on vo I^s- sμ m 00 o vo on H m sμ sμ CM ro CM CM o o o cn o ro sμ ro sμ sμ in VD vo r- 00 I^s- vo in in in m in m m in sμ in m m in in in m in m m m m

co en cn en cn cn cn en en cn en cn o o o o

fit fit <-ϋ fit fiϋ fiϋ fit fiϋ ft fiϋ fiϋ fit ft ft fit fit f fi fit fiϋ fit to 0 0 0 0 0 0 0 0 0 0 0 ft fit fit fit o o O O O Pi Pi Pi Pi Pi Pi Pi Pi Pi P I Pi Pi Pi P u f ft fiϋ ft ft fiϋ fit fiϋ ft fit fiϋ fit fi fiϋ fit fiU ft ft ft ft ft

CM fit PQ CD P PQ N X PQ fit PQ ft PQ CD P

O a U U U U a U a a U O a U U u O a U U U U cn o CM sμ I^s- o ro in vo en CM ro sμ vo co CM ro sμ m I^s- o ro en o O o o H H H H H CM CM CM CM CM ro ro ro ro ro sμ s sμ m in m in m m m m m m m in m m m m m in m m m

CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 O O O O O O O O O O O O O O O O O O O O O O EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH ι=l^' <i ⁽ ι^ f^ ^ ⁽ f f ; ι ; f =t^" ι^ ι<C ^ fi^ fi^ fi^ '=3l fi^:i; u o a u u u u a u a a u o a u u u o o u o a

I^s- CM oo I- v- cn sμ o m oo ro o H o in sμ I^s- in CM I^s- sμ ro

I- o oo rH irI- vo tr- sμ H oo vo CM m rH -i ro o I^s- co ro on oo co oo on on H sμ r- o H o oo co co co oo VD s CM I^s- r- I^s- o o o o o v- rH H CM CM CM o o o o o o o O o o o rH H rH H H v-i H H H H H H H H H H H H H v-i H v-

O O O o o o o o o O o o o o O o o O O o O σ o O o o o o o o o O o o o o O o o O o o o o

H H rH H H H v-i H v-i H H H H H H H H H H H H v-i

VD sμ m on cn co o I^s- on ro ro m CO on on I^s- ro O CM sμ 00 cn

O ro sμ in r- H on vo sμ CM vo CM in cn sμ vo on I^s- H in ro m sμ CM sμ o H VD m ro sμ VO ro oo o sμ ro CM H O ro in H H sμ sμ ro CM H rH o o cn CO on H H CM CM ro ro sμ CM CM CM CO

CM CM CM CM CM CM CM CM v-i H H CM CM CM CM CM CM CM CM CM CM CM 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 en rH CM cn o VD o cn CM on oo CM ro co oo ro H I^s- cn r- H cn m ro ro in co H H cn in ro on -i H r- m H VD H sμ Oϊ cn H

VD I^s- cn ro CM CO cn vo r- co vo I^s- in cn CM ro VD in cn sμ m ro ro sμ CM ro CM H o on oo co I^s- sμ m sμ VD VD I^s- ∞ I^s- r- ∞ I^s- in in in m m in m sμ sμ sμ sμ m in m in ω ιn in m in in in o in CM CM o cn m m o r- m sμ o ro o H CM ro ro ro m oo co CM CM CM vo m ro sμ o m I^s- en I^s- CM irCM oo vo H r- ro on co ro sμ sμ o sμ H ro o vo I^s- o ιn CM on CM cn I^s- CO o s oo r- r- co oo on o rH o o H cn cn oo o o CM CM CM ro o o co m m in m in vo vo vo o vo m in m VD vo VD vo VD vo vo vo m

rH v-i CM CM CM CM CM CM CM CM CM CM CM ro ro ro ro ro ro ro ro s

CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM

CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM ft fit fit ft fit fit fit fit fit fit fit ft fit fit fit fit ft fit fit fit fit fit o o ø ø ø ø ø ø ø ø ø ø ø ft ft ft ft ft ft ft ft p Pi P^ p-l p-j p-i Pi P-J p-j p-l p-J p-l P-J ptJ to co to to to to ω to pQ ft ft I ι <; ι ι f f ι <; ; ι rt' f ι ι f-⁽; rf! rQ rH CM H CM fit, PQ CD P PQ N K X fit PQ CD P P u O a U U U U a u a a υ O a U U υ O O U O a

VD I^s- oo o CM m oo H co sμ I^s- o H CM sμ vo cn o H CM ro sμ sμ sμ sμ in m L0 in VD vo VD VD I^s- I- I^s- ir- i-- r- 00 CO oo oo oo in m in m m m in in m m m in m m LT) in m m m m m in

CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 O O O O O O O O O O O O O O O O O O O O O O EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH fi fi fi fi fi fi fiϋ fi fiϋ fi fi fiϋ fiϋ fi fiϋ fiϋ fi fiϋ fiϋ fiϋ fi fi ATOM 2586 CA LEU A 224 57..836 58..325 -23..068 1..00107..84 c

ATOM 2588 CB LEU A 224 56. .775 58. .140 -24. .181 1. .00107. .59 c

ATOM 2591 CG LEU A 224 57. .160 57, .918 -25. .674 1. .00107. .02 c

ATOM 2593 CD1 LEU A 224 55. .922 57. .933 -26. .552 1. .00106. .52 c

ATOM 2597 CD2 LEU A 224 58. .158 58. .894 -26. .280 1. .00106. .57 c

ATOM 2601 C LEU A 224 57. .166 58. .385 -21. .664 1. .00108. .13 c

ATOM 2602 O LEU A 224 55. .968 58. .494 -21. .610 1. .00107. .71 0

ATOM 2603 N CYS A 225 57. .941 58. .301 -20. .559 1. .00108. . 69 N

ATOM 2605 CA CYS A 225 57. .579 58. .889 -19. .230 1. .00108, .83 C

ATOM 2607 CB CYS A 225 58. .590 58. .569 -18. .132 1. .00108, .56 C t o ATOM 2610 SG CYS A 225 59, .239 56, .908 -18. .039 1. .00107, .86 s

ATOM 2611 C CYS A 225 57. .605 60. .415 -19. .273 1. .00109. .57 c

ATOM 2612 O CYS A 225 56. .901 61. .089 -18. .537 1. .00109. .18 0

ATOM 2613 N GLU A 226 58, .517 60. .940 -20. .082 1. .00110. .74 N

ATOM 2615 CA GLU A 226 58. .595 62. .358 -20. .369 1. .00111. .54 C

ATOM 2617 CB GLU A 226 59. .839 62. .645 -21. .230 1. ,00111. .94 C

ATOM 2620 CG GLU A 226 60. .290 64. .103 -21. .276 1. .00112. .83 C

ATOM 2623 CD GLU A 226 59. .796 64. .803 -22. .523 1. .00114. .49 C

ATOM 2624 OE1 GLU A 226 59. .888 64. .227 -23. .632 1. .00115. .97 O

ATOM 2625 OE2 GLU A 226 59. .295 65. .929 -22. .394 1. .00115. .47 O

ATOM 2626 C GLU A 226 57. .311 62. .717 -21. .089 1. .00111. .81 c

ATOM 2627 O GLU A 226 56. .388 63. .277 -20. .494 1. .00111. .40 o

a u u u u a u a a u o a u u u to u u o a u u

I^s- I^s- cn CM co o sμ m o CM cn sμ cn oo ro o cn CM cn vo r-i vo I^s- vo 00 VO oo CM ro CM ro in H ro o o cn cn en in ro vo o o

CM ro ro sμ vo co cn o o ro ro ro ro ro CM CM H CM H CM ro ro H rH H H H H H CM CM H H H H H H H H H H rH r-i H H rH H H rH v- H H H rH H r-t H H rH H H v-i H H rH H O O o o O O o O o o o o o o O O o o o O O o O O o o O O o o o o o o o o O O o o σ O O o

H rH H H v-i H H H H H H H H rH H H H H H H H H ro 00 in H o CM vo cn VD m CM co VD VD sμ ro sμ H in H sμ O m CM CM sμ on I^s- oo H I^s- D H on sμ sμ m in cn ∞ on CM H sμ ro CM o co o en in ro sμ m O sμ in co vo on o in en ro ro on

CM ro sμ in vo vo o m CM ro rH o on o cn v-i en cn oo I^s- m

CM CM CM CM CM CM CM CM CM CM CM CM CM rH CM rH CM H -i rH rH rH 1 1 1 1 1 1 1 1 I 1 1 1 1 1 1 1 1 1 1 1

I^s- rH ro 00 ∞ co vo ^■sμ H H CM vo rH cn in CM H VD ro cn sμ o O in I- ro VD ro VD m co I^s- O I^s- on in r- CM oo σ sμ t- ro ro in oo vo I^s- vo sμ O LΠ on CM CM en m ro o cn H ro cn cn ro

CM CM H CM CM H o O cn -i CM rH o on oo vo CM ro H CM CM vo VD VD VD vo VD vo VD m vo VD VD vo m in m in VO VD vo vo vo

H m co oo I^s- CO CM in cn H sμ I^s- I^s- o oo CM in CM I^s- sμ CM I- sμ m H o 00 O oo en sμ ro o cn CM ∞ CM co o CO VD H CM I-

CM o ro on ro cn m VD H r- vo oo oo cn sμ H CO VD CM CM CM

I^s- vo vo m sμ ro ro ro ro sμ ro sμ ro ro ro ro m ro ro sμ sμ sμ m m in in in m m m m m m m m in in m m m m m m m

t- I^s- I^s- I- I^s- I^s- I^s- I^s- I^s- I- t- oo oo oo oo oo oo oo oo cn cn cn

CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM

CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM ft fit fit fit fit ft fit f fit ft fit fit fit f ft fit fit fiϋ fit fit fit

0 0 0 0 0 0 0 0 0 0 ^ EH EH EH EH EH EH EH EH fit fit fit

P Pi Pi Pi Pi Pi Pi Pi Pi fYj PQ PQ PQ PQ PQ PQ PQ PQ μQ μQ μQ fit fit fiϋ fit ^ fiϋ ft <; fit ft 2 2 2 2 2 2 2 2 fϋ fiϋ fit CM a fit PQ 0 P PQ IS] X K f PQ 0 P PQ f PQ

U U U u a u a a u O a U U U to U U O a U U

∞ o CM in ∞ H co s]< I^s- o H CM sμ vo on CM ro r- ∞ cn rH ro

CM ro ro ro ro sμ sμ sμ sμ m m m m m m vo vo VD vo vo I^s- I^s-

VO vo vo VD vo vo vo VD vo VD VD vo vo vo VD vo vo vo VD vo VO vo

CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM

2o 2o 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2

O O O O O o O O O O O O O O O O O O o O

EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH fit fit fit fit f fit fit fit ft fiϋ fiϋ f fit fit ft fit fit fit fit fit fit ft u o a u u u o a u u u o a u u u u u u o a u

vo o I^s- H o o m co on m m sμ CM CM cn co cn sμ vo I^s- on v- sμ rH CM o ro CM sμ co CM H cn H in o H o I^s- oo sμ ∞ co σ ro ro sμ sμ sμ m in m VD vo vo I^s- I^s- oo ro en en on oo I^s- cn o v-i H H H H H H H H H H H H H H H H H H H H CM H rH H H H v- H H H H H H H H H H H rH H H H H o O O o o o o o O o o O o O O σ o O o O O O o o O o o o σ o O o o o o O O σ o O o O O o

H rH H H H H H H H H H H v-i H H H H rH H v-i H H

I^s- in ro on sμ oo ro in ∞ co VD in sμ ro H VD I^s- cn CM O H cn o m sμ sμ o in σ 00 I^s- sμ σn VD sμ ΓO ro in ro VO ro oo m I^s- m I^s- in in oo sμ 00 on on ro co oo sμ I- VD o on ro CM sμ m

I^s- VD 00 ∞ on oo ∞ I^s- r- I^s- VD in in sμ ro sμ sμ sμ sμ ro m m

H H H H H H H H H H H H H -i v-i H H r-i H H H H

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 m H sμ oo cn t- I^s- ro r- CM sμ H vo VD co sμ vo CM vo I^s- co

CM H ro I^s- ro cn m VD H co r- o o O sμ vo sμ CM co vo μ in o H oo H sμ o CM in ro ro on CM ro r- I^s- t- CM vo H m co CM sμ sμ sμ D vo I- co vo r- vo I^s- oo oo co r- VD vo in o o o CM vo vo VD vo vo VD vo VD vo vo VD VD VD VD vo VD vo VD I^s- I^s- t- I^s-

I^s- o I^s- CM r- o co CM ro I^s- VD CM ro cn ro oo I^s- o VD ro CM ro sμ CM oo ro o CM H oo sμ CM on O t- sμ o CM CM co ∞ oo m CM ro ro v-i I^s- in in VD ro v-i cn o o CM sμ sμ H sμ H σn sμ co CM in VD m ιn vo sμ sμ ro CM o CM rH ro CO sμ m vo sμ co sμ ro sμ in in m in in in in m m m in m m in in m m in in in in in

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PQ PQ PQ ft ft ft ft ft ft ft ft PQ PQ PQ PQ PQ PQ PQ PQ P P P

K tn K to CO to to co CO to co P μQ l-l μQ μQ μQ μQ μQ μQ μQ μQ ft ft ft fit fiϋ fit fit f ft fit fiϋ H H H H H H H H 0 0 0

CM H CM H H CM

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2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 o EH oEH oEH oEH oEH oH oEH oEH oEH oEH oEH oEH oEH oEH oEH oEH oEH oEH oEH oEH oEH oEH ft fit fit fit fit fit fit fit fit fit ft ft fit fit fit fit ft fit fit fit fit fit u u o o u o a u u to u o a u u u u u a u o a

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o o o σ o o H H H H vi vi CM CM CM CM CM CM CM CM CM ro tr- c- irt- I^s- c- I irI^s- I^s- I^s- irI^s- t- c- r- I^s- irirt c- I^s- ro ro on ro ro ro ro on ro ro ro on ro ro ro ro ro on on cn ro ro f fit ft fit fit f fit fit fit fit ft ft f fit fit fit fit fit f fit fit f

P P P P P P to to to co o to co O CO CO CO co CO co CO f μQ μQ μQ μQ μQ μQ i H H !H !• H H H H H >H H t>H μQ

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2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 O O O O O O O O O O O O O O O O O O O O O O EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH ft ft ft ft fit fit fit ft fit fit fit fit fit fit fit fit fit fit fit fit fit fit u u u o a u u o a u u o a u u u u o o u o a

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H H rH rH vi vi vi rH H rH vi vi H vi vi H H H H H r-i H s ro O oo on CM co co CM vo cn r- ro CM CO I^s- co CM CM vo IT- in

H cn ro vo I^s- 00 cn o I^s- on cn o o VD ro o o sμ H in CO in vo vo CM cn sμ CJ sμ m cn m r- oo m ro sμ en co o H sμ VO on m in VD I^s- in in in m sμ sμ m vo in VO m sμ ro CM sμ t- CO VD ro ro co ro cn cn ro ro CO ro ro cn ro ro ro ro ro ro ro ro ro ro

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VD H Γ- o o CM ro CM cn CO sμ sμ in o vion tr- o co in ro CM cn in CM I^s- H VD on m vo ∞ on o o on vo CM CM vo H in vo H σ cn H v-i rH o rH on oo iroo in sμ sμ co on CM in m in sμ ro sμ sμ sμ sμ sμ sμ n ro on n ro ro ro cn cn cn cn cn ro ro

ro cn cn cn sμ sμ sμ sμ m m m if) VD vo VD VD VD VD VD D vo I^s-

I^s- I^s- I^s- I^s- I^s- irr- I- I^s- I^s- I^s- I^s- irr- I^s- I^s- ro ro ro ro on ro on ΓO ro ro ro ro ro on on on on o n ro ro ro ft fit fit fit fit ft ft ft f f fit fit fit fit fit fit rti fit fiϋ ft fit fiϋ f fit fit fit H H ^H H !» H >i H P P P P P P P P P ft μQ μQ μQ μQ μQ μQ μQ μQ μQ μQ P P P μQ μQ μQ μQ μQ μQ μQ μQ to fit fit ft fit 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 fit rH CM ft PQ f fiϋ fiϋ ffl 0 P PQ PQ

U U U O a U U b a U U b a U U U U b b U O a

CM sμ oo on o CM m vo I^s- cn CM ro sμ vo oo H sμ m VD r- 00 on co co oo oo on on cn on cn cn o o o o o rH H rH H rH rH i t- t- I- I^s- I^s- I^s- t- ir- I^s- t- oo oo oo oo oo 00 co co oo oo CO oo sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 o b b b b b b b b b b b b b b b b b b b O O

EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH fit fit fit fit fit fit ft fiϋ fiϋ ft ft fiϋ fiϋ fit fit fit fit fit fiϋ fit ft ft

111 u u u o o u o a u u u u o o u o a u u u u u

ro cn CM CM vi ΓO cn r- in m ro I^s- CM m o o vo ∞ in I^s- ro r- o VD co VD VD ΓO on H on vo on vi 00 vi sμ o vo co ro m cn sμ sμ VD cn vo VD I^s- o co vi H o m vo ro CM CM sμ D sμ in in in m in m m in in m vo vo I^s- m in in in m m in in o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o σ o o o o rH H vi -i vi H H H -i H H H H rH H rH vi H H rH vi H

VD CM cn I^s- ro sμ o O irsμ o oo CO VD co CM vi co 00 I- co VO vo sμ oo m irI^s- VD VO on irsμ VD sμ cn t- sμ cn CM cn m VD oo

I^s- CM on VD on I^s- co co 00 on VD CM sμ m o cn on σ H m m cn

I^s- I^s- I^s- I^s- oo I^s- vo oo CO cn on O H cn cn cn oo cn o rH CM H ro ro ΓO ro n ro ro ro cn ro ro sμ sμ cn ro ro cn cn sμ sμ sμ sμ

H vo CM sμ o CM oo r- VD H oo H oo I^s- oo t— n cn o cn in cn 00 m H oo vi sμ o VD vo CM o co vo CM vi vo vi r- in CM vi oo o sμ CM m co cn v-i oo cn sμ irVD CM o If) r- vi VD sμ cn ro VD sμ ro CM vi H CM CM CM H CM on cn ro sμ o cn o oo oo oo CO oo co oo oo oo 00 co oo oo oo 00 00 oo co co 00 r- oo I- I^s- I^s- I^s- r- o H ∞ o ro cn I^s- vo r- I^s- on oo CM on o ΓO CM m CM co ro tn

CM H H I^s- ro sμ CM co cn I^s- in sμ 00 I^s- m vo oo vo o CM on I^s- vo on on oo vo vo I^s- cn sμ sμ CM vo vo vo vi o oo sμ sμ I^s- t- H sμ ro CM vi ro in m vo t- oo cn o o r- oo m m sμ sμ ro VD co ro cn cn ro cn ro ro ΓO ro ro sμ sμ sμ m ro cn ro CO ro ro ro

I^s- r- I^s- I^s- I- r- I^s- co oo oo oo oo co co co oo on cn cn cn cn cn

I^s- I^s- I^s- 1- I- I^s- I^s- I^s- t- irirI^s- r- r- r- I^s- I^s- I^s- r- I- c- r- ro ro ro ro ro ro ro ro 00 on on ro ro ro ro ro cn ro ro ro ro ro

rf; f ι f ( rf; f-i! rf; f- fiϋ fiϋ ft ft ft ft ft _ft _ft p p p p p p p p p p p p p p p CO tΛ tO tO tO CO CO P μQ P P P P P PQ pQ pQ pQ pQ pQ fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ ø ø ø ø ø ø ø ø ø μ^Q P ^μQ P ^μQ μ^Q

H cn vo I^s- oo on o H ro in co H CM ro sμ in vo 00 o cn m cn

CM CM CM CM CM CM ro cn ro ro 00 sμ sμ sμ sμ sμ sμ s in If) in in

CO oo ∞ oo oo 00 ∞ oo oo ∞ oo 00 ∞ oo co oo oo oo ∞ ∞ co oo sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ μ

2o 2o 2o 2o 2o 2b 2b b2 2b 2b 2b 2b 2b 2b 2b b2 2b o2 o2 2o 2o 2o

EH EH EH EH EH EH R EH EH EH EH H EH EH EH EH EH EH EH EH EH H u b a u u u b a u u u u u u u u u u a u u u

o sμ on m cn oo in oo cn o VD sμ CM en vi H sμ I^s- cn H cn in in I^s- I^s- CM sμ H CM rH cn H H oo m oo VD ro ro vo ro cn H t--

CM ro vi o o oo vo I^s- vo 00 in r- o CM H o ΓO m o oo co oo in m m in m sμ sμ sμ sμ sμ in m vo vo vo VD sμ sμ sμ cn ro ro o o o o o o o o o o o o o o o o o o o o o O o o o o o o o o o o o o o o o o o o o o o O

H vi H rH rH H H rH v-i vi H H vi vi H vi H H vi i vi oo vo I^s- r- CM sμ CM I^s- If) if) VD vi H vi H on CD o VD if) CD r-

CM m O rH vo H CM cn in CM VD I^s- cn o o CM CM v-i m vo I^s- o

I^s- vo I^s- ro ro on ro o in VD ro cn vi on cn H 00 cn cn CM cn r-

I- I^s- vo m sj< sμ m sμ ro CM CM cn cn vi o H CM CM CM CM CM ro ro ro ro ro ro ro ro ro ro ro ro cn cn cn 00 ro CO ro ro ro ro 00 oo I^s- oo rH o o H CM co o c- co vo vo CM in rH CM CM vo CO sμ cn o cn cn CM ro sμ oo rH CM CM co i^rCM VD VD sμ I^s- sμ sμ cn H o on cn sμ I^s- in VD in vo vo sμ sμ on CM H CM sμ sμ H on in sμ oo VD ∞ co cn I^s- I^s- vo in sμ ro CM H H CM ro VD I^s- vo vo vo m iT- I- I^s- I^s- I^s- I^s- I^s- r- r- r- I- I^s- I^s- I^s- I^s- I^s- I^s- I^s- I- r- I^s- r- cD sμ VD m CM ir- oo VD m CM o co I^s- ro I^s- cn m CM I^s- o r- I^s- o o O sμ sμ sμ o m H sμ oo oo cn CM sμ cn vi I^s- I^s- co I^s- sμ cn m co o vo r- cn cn cn D sμ vo sμ H cn rH cn cn in m oo vo sμ sμ sμ sμ sμ ιn vo if) VD VD VD I^s- co I^s- r- irVD co cn o o cn ro ro n ro ro ro cn ro ΓO ro ro ro cn ro ro on ro ro ro sμ sμ

cn on o o o o o vi H H i vi vi vi H v-i H H CM CM CM CM t- irco co co co CO ∞ oo oo co co oo oo co co oo oo oo co oo oo ro on ro cn cn 00 ro ro on cn cn ro ro cn cn ro cn cn co ro cn cn fit ft f f fit fit fit fiU fit fit f fit ft fit fit fit f fit fit fit fit

P P fit fit fit fit ft, PQ PQ PQ PQ PQ PQ PQ PQ PQ PQ PQ O O O O

PQ PQ μQ μQ μQ μQ μQ PQ X K X X K ffi PQ X K X Pi Pi Pi Pi μQ μQ ft fit f ft ft ft ft ft ft ft ft ft ft ft ft ft ft ft ft ft

H vi CM t Pi fiϋ PQ CD P PQ IS] PQ P fit ffl CD

U b a υ υ U O a U U U U U U U U U O a U U U ro sμ in r- on n sμ in r- cn CM ro m cn H ro sμ m VD co H

VD vo vo VD vo ir- r- r- I^s- oo co co co co on cn on cn cn σn o oo co ∞ co co co CO co oo oo oo oo oo co oo co co co ∞ co oo cn sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ S sμ sμ sμ sμ sμ sμ sμ sμ sμ

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 O O O O O O O O O O O O O O O O O O O O O O EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH ft fit fit fit fit fit fit ft fit fit fit fit fit fit ft fit fit fit fit ft fit fit ATOM 4904 CD PRO A 382 39..184 75..158 33.869 1..00 40,.24 c

ATOM 4907 C PRO A 382 39. .629 76. .690 30 .740 1. .00 37, .52 c

ATOM 4908 O PRO A 382 39. .201 75. .677 30. .234 1. .00 35, .77 0

ATOM 4909 N VAL A 383 40, .102 77, .693 30, .026 1. .00 37, .08 N

ATOM 4911 CA VAL A 383 40. .131 77, .650 28, .593 1. .00 37, .10 C

ATOM 4913 CB VAL A 383 38. .905 78, .476 28. .063 1, .00 38, .33 C

ATOM 4915 CGI VAL A 383 38, .909 78 .606 26 .522 1. .00 3 .29 C

ATOM 4919 CG2 VAL A 383 37, .578 77, .875 28 .602 1. .00 34, .93 C

ATOM 4923 C VAL A 383 41. .453 78. .221 28, .081 1. .00 36. .74 C

ATOM 4924 O VAL A 383 41. .778 79. .362 28. .332 1. .00 36. .52 O

ATOM 4925 N ALA A 384 42. .187 77. .373 27. .378 1. ,00 36. .82 N

ATOM 4927 CA ALA A 384 43. .506 77. .673 26, .844 1. .00 37. .74 C

ATOM 4929 CB ALA A 384 44, .026 76. .513 26, .142 1. .00 36. .77 C

ATOM 4933 C ALA A 384 43, .578 78. .901 25, .926 1. .00 39. .73 C

ATOM 4934 O ALA A 384 44. .593 79. .634 25, .960 1. .00 42. .24 O

ATOM 4935 N GLY A 385 42. .537 79. .170 25. .142 1. .00 40, .49 N

ATOM 4937 CA GLY A 385 42. .465 80. .420 24. .376 1. .00 40. .85 C

ATOM 4940 C GLY A 385 42. .558 81. .680 25, .205 1. .00 41. .33 C

ATOM 4941 O GLY A 385 42. .923 82. .752 24. .738 1. .00 39. .42 O

ATOM 4942 N HIS A 386 42. .175 81. .535 26. .457 1. ,00 43. .76 N

ATOM 4944 CA HIS A 386 42. .195 82. .644 27. .379 1. ,00 46. .19 C

ATOM 4946 CB HIS A 386 41. .294 82. .410 28. .597 1. ,00 47. .15 C

u a u a u u o a u u u u u o a u u u u o o u

cn CO o H o CM sμ cn co co sμ CM m co H CM sμ I^s- co I^s- oo irsμ sμ cn co o cn in cn CM o n vi o cn CM I- o m sμ m oo on cn CM CM H H in vo in vo vo m VD I- oo cn o H H m sμ cn sμ in in m in sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ m in m sμ in sμ o o o o o o o o o o o o o o o o o o σ o o o o o o o o o o o o o o o o o o o o o σ o o o

H vi vi i vi rH H vi H H vi H H H H vi rH H vi vi H i cn cn cn VD in cn in I^s- H VO ir- VD H I^s- in vo m sμ sμ vo σn I^s- en oo cn CM IT- I^s- H sμ O r- cD sμ O m co on I^s- o vi vi CM cn

CM o H ro CD oo sμ vo CM VO o I^s- oo ro sμ on vo on co H o m oo I^s- r- co cn I^s- oo I^s- CO I^s- I^s- VD cn o o vi CM H σ H on CM

CM CM CM CM CM CM CM CM CM CM CM CM CM ro n cn cn ro cn on CM n o H en o I^s- m sμ m O co sμ O sμ o if) co ro CM CD o I^s- CD ro ir- in oo vo in o oo 00 I^s- CM VD o cn CM I^s- m CM D I^s- CM r- ro co oo CM m r- oo co H in H on CM cn sμ ro CM sμ o sμ CM cn

CM H H CM CM CM H cn sμ if) VD sμ sμ ro sμ sμ m vo vo in VD CM oo oo oo oo oo ∞ co co 00 ∞ 00 co co 00 oo oo oo co CO ∞ co oo

VD oo o oo CM sμ m cn cn r- CM 00 o H r- cn in sμ vi o o vo ro sμ ro sμ VD o on sμ CM ∞ cn on vi I^s- cn CM m H H CM I- cn 00 cn o VD ir- VD o CM m oo VD r- in in ro sμ cn I^s- I^s- vo o CM cn on oo i^rco cn sμ sμ m if) sμ ro in vo sμ sμ cn CM i o CM sμ co cn ro on ro sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ s}< sμ sμ

vo VD vo vo VD vo vo I^s- I^s- I^s- r- I^s- r- I- oo oo co 00 co oo oo oo oo 00 ∞ oo co oo oo co oo co oo oo oo co oo oo co CO CO oo oo oo cn ro ro ro ro ro ro ro ro ro ro on ro n cn ro ro ro ro ro ro n ft f fit fit ft fit fit ft fit fit fit f f f fit fit ft fit ft ft fit fit co to to to CO CO to O O O O O O O P P P P P P P P

H H l-H H H H H Pi Pi Pi Pi Pi i Pi μQ μQ μQ μQ μQ μQ μQ μQ

K X X X K X ft ft ft ft ft ft ft 0 0 0 0 0 0 0 0 H H CM CM H CM

CD P PQ PQ P ft PQ CD P ft ffl 0 P PQ PQ

U a U a U U o a υ υ U υ υ O a υ υ U U υ O υ cn o CM sμ VO ∞ on o rH ro VD on CM ro sμ VD oo H sμ in vo I- sμ m m in m in m vo VD vo VD VD I^s- i^rirl— I^s- CO co 00 oo oo en cn cn cn on cn on cn on on CD on cn o on on en on on cn cn on sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 O O O O O O O O O O O O O O O O O O O b b b EH EH EH H EH EH EH EH EH EH E-J EH EH EH EH EH EH EH EH EH EH EH ft fit fit ft fit fit fit fit fit fit ft fit fit fit fit ft fit fit fit fit fit fit o a u u u o o u o a u u u u u u o a u u u u

H r- in in H CM ro H oo o m I^s- cn o t- o oo H ro ro in CM sμ sμ H m vi cn en m vi oo 00 cn VD co oo cn t- CM CM sμ o oo o cn o on 00 cn VD o cn o o o co m sμ o o vi o o o oo m sμ in sμ sμ sμ sμ m in in m m m in in m in if) in m m sμ o o o σ o o o o CD o o o o o o o o o CD o o o o o o σ o o o o CD o o o o o o o o o o o o o rH H rH H rH rH rH rH vi rH vi H rH rH H vi H rH -i H vi rH

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CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM ro cn on CM CM

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ro ro ro ro co ro ro sμ sμ sμ sμ sμ sμ sμ sμ m in m m in m in

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U u U υ U υ o a U U U o O U O a U υ U to υ U o CM sμ t- vi m vo IT- on vi sμ m VD CO on i cn vo CD o sμ

I^s- r- irI- oo oo oo CO co CD on on on on on on o CD o CD rH r-i cn cn on σn on cn on cn σn on on on on cn on on o CD o o o CD

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m VD VO vo vo VD vo VD vo VD vo VD VD I- I^s- I^s- I^s- co oo oo co ∞

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H Pi Pi Pi Pi Pi Pi Pi Pi Pi Pi Pi Pi !χ tx tx !χ PQ PQ PQ PQ PQ

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CM ro sμ m CD sμ o VD H ∞ m vo I^s- in on sμ cn vi cn vi o VD oo on σ m sμ sμ co in on ro sμ 00 CM VD in VD CM I^s- en o in sμ on o σ H vi CM vi CM CM vi o CM CM ro sμ ro H rH cn cn cn on sμ m in m m m if) in m if) in in in m m m m m sμ sμ sμ sμ vo H cn o on o I^s- oo CM VD VD cn CM VD oo vo H VD o o co H oo oo VD vi H vo oo in o co sμ cn sμ o cn ro co in rH i cn r- m I^s- cn oo I^s- oo VD on m CM m 00 vo CM H I^s- I^s- r- en sμ I^s- t- sμ H σ i o o H CM cn cn co oo I^s- r- I- in I^s- I- IT- oo cn on sμ sμ s sμ sμ sμ sμ sμ ro ro cn ro ro oo ro ro ro ro CO cn on ro

co oo 00 cn cn on on σn cn on σn o o o o o o o vi H H vi

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2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 O O O O O O O O O O O O O O O O O O O O O O EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH ft ft fit fit ft fit ft fit fit fit ft ft fit fit fit fit fit ft ft fit fit fit o o u o a u u u u u a u o a u u u u u u o a

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H H H vi H rH H H H vi H H H H vi H H H

CD o o o o σ σ σ σ σ o O σ o O O O o o O O o

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00 cn VD in t- oo cn oo oo I^s- VD on on o vi cn ro sμ ro rH o CM

CM CM CM CM CM CM CM CM CM CM CM CM CM ro cn cn ro ro ro ro co cn

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rH rH H rH CM CM CM CM CM CM CM CM CM ro ro ro ro oo ro ro ro sμ ro ro ro ro ro ro ro ro oo oo oo ro ro ro ro ro ro ro ro ΓO ro ro Ln Ln m in in in in in in m in in in in in m m m m m m m ft fit fit fit fit fit fit ft ft fit fit fit fit fit fit ft ft fit fit fit fit fit ft CLi Ci ft to c c to CO CO to CO C0 PQ Pϋ PQ pq pQ PQ pq p-i to co to to co tx t !χ H tx tx tx !^χ !χ μQ P P μ-l μQ μQ μQ μQ I-H f ft f f μQ μQ μQ μQ μQ μQ μQ μQ μQ H H H H H H H H p rH CM H H CM

P P rfl ffl U P PQ -o o o u o a u u u u u a u o fiut fflu 0u Pu 0u u o a sμ in VD I^s- ∞ o CM ιn ∞ H sμ co cn O CM sμ VD cn ro I^s- oo on o o O σ o H H H vi CM CM CM CM cn ro cn ro ro sμ sμ sμ sμ

H H H H i H H H vi H rH H rH vi vi r-i H H rH vi H H

I^s- t- I- t- t- c- C- I^s- I^s- tr- ir- C- I^s- I^s- I^s- I^s- I^s- I^s- r- I^s- irI^s-

o 2 2o o2 o2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2

EH EH EH EH oEH oEH oEH oEH oEH oEH oEH oEH oEH oEH oEH oEH oEH oEH oEH oEH oEH oEH ft fit fit fit ft fit fit ft fit fit ft fit ft fit fit fit fit fit fit fit fit tt ATOM 7151 CA LYS A 534 33..435 48..205 32..252 1..00109..40 c

ATOM 7153 CB LYS A 534 32. .214 47. .311 31. .910 1. ,00109. .47 c

ATOM 7156 CG LYS A 534 31, .889 47, .231 30. .387 1. .00110. .28 c

ATOM 7159 CD LYS A 534 31, .361 48, .570 29. .789 1. .00110, .30 c

ATOM 7162 CE LYS A 534 31. .383 48. .590 28. .251 1. .00110, .62 c

ATOM 7165 NZ LYS A 534 32, .318 49, .631 27. .700 1. .00111, .07 N

ATOM 7169 C LYS A 534 33, .187 49, .084 33 .553 1. .00110, .07 C

ATOM 7170 O LYS A 534 33 .789 48, .836 34. .642 1. .00109, .96 O

ATOM 7171 N LEU A 535 32. .339 50. .119 33. .427 1. .00109. ,98 N

ATOM 7173 CA LEU A 535 32. .085 51. .017 34. .559 1. .00110. .21 C

ATOM 7175 CB LEU A 535 33, .381 51. .735 35, .000 1. .00110. .14 C

ATOM 7178 CG LEU A 535 34. .048 51. .252 36. .293 1. .00110. .93 C

ATOM 7180 GDI LEU A 535 35, .574 51, .220 36. .148 1. .00110. .56 C

ATOM 7184 CD2 LEU A 535 33. .615 52, .079 37. .546 1. .00112, .29 C

ATOM 7188 C LEU A 535 30. .986 52, .060 3 .326 1, .00110, .22 C

ATOM 7189 O LEU A 535 30. .500 52, .273 33. .206 1. .00110, .17 O

ATOM 7190 N SER A 536 30. .602 52. .690 35. .434 1. .00110. .12 N

ATOM 7192 CA SER A 536 29. .640 53, .785 35, .440 1. .00109. .79 C

ATOM 7194 CB SER A 536 28, .562 53, .541 36, .515 1. .00109. .90 C

ATOM 7197 OG SER A 536 27. .594 52. .593 36. .061 1. .00110. .16 O

ATOM 7199 C SER A 536 30. .355 55, .114 35. .662 1. .00108. .92 C

ATOM 7200 O SER A 536 29. .738 56, .168 35. .524 1. .00108. .80 O

a u u o u o a u u u u u u o u u u o a u u u

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H vi H vi H vi vi H H H H H H vi rH H H rH H H o CD O o o o O o O O O o o CD o o O O o o o o o CD O o o o O o o O o o o CD o o o O o o o o

H v-l vi H H H vi vi H H v-i H vi i H rH H H H H H H in vi CM m sμ VD sμ vi in O CM CM CD if) H r- H cn O CM 00 O

CM CD CO sμ CM ro o CM o H CO VD vo CD sμ cn I^s- co sμ H co o vi sμ vo oo m oo sμ on 00 VD O H VD co ro 00 CM CO m VD o vo VO t- r- in vo sμ sμ CM CM H H H O H CM sμ sμ in vo r- 00 ro ro ro ro on ro ro ro ro ro ro ro on ro rO ro ro cn ro ro ro cn r- o ro in vi sμ vi cn CM I^s- I^s- vo sμ co cn co vi vi vo 00 vo in sμ CM on VD o CM cn CM CM CM m ro oo on ro m m H sμ VD o 1- o CM oo 00 co cn sμ CM H on o on VD m m D ro in H o on CM m VO VD I- m sμ VD vo VD sμ L0 ro CM vi CM ro I- I^s- oo on on H m in m in m in in m in in m m m in m m in m m m m vo o ro vi o ro i m cn sμ on I- o in o cn en oo CM o cn CM CM sμ ro I^s- o oo i on o CO VD in ro CO rH sμ m m CM vo I^s- in H

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2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 O O O O O O O O O O O O O O O O O O O O O O EH EH EH H H EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH ft fit fit fit fit ft fit fit fit ft fit fit fit fit fit ft fit fit ft fit fit fit u u a u o a u u u u u u o a u u u o a u o a

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CM CM CM ∞ t- oo 00 cn 00 t- cn I^s- vo 00 I^s- oo o H o vo m VD vo o vo in m m in m m in in in m m m in VD vo VD in m in m sμ on o in ∞ sμ CM in CM in co sμ sμ r- on CM co m D sμ oo vi co H CM CM vi H sμ o ro CO in t- vi CM σ H CM VD CM irm m sμ o VD in i^rcn vi in in in r- sμ oo VD oo cn H oo cn on co irvo in oo co on o CM ro ro sμ o rH on on co oo 00 cn 00 cn t- on ro ro cn ro ro sμ sμ sμ sμ sμ sμ sμ ro ro cn ro cn on ro ro ro

cn en cn on cn O o CD CD O o o o H H H H H H H H CM cn cn ro ro ro sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ s sμ sμ sμ sμ sμ sμ in if) m in m in in in in m in m m m m in in m m m m in fit fit ft fit fit fit fit fit ft fit fit fit fit fit fit fit fit ft ft fit fit fit to to to to to P P P P P P P P a a a a a a a a fit tx tx tx tx tx PQ PQ PQ PQ PQ PQ PQ PQ co co O to co O co co μQ μQ μQ μQ μQ μQ μQ μQ i-l μQ μQ μQ fit fit ft f fit fit fit fit fit H

P PQ ISl fit ffl CD P P fit ffl CD P P u U a U o a U U u υ U O a υ 0 υ υ a υ o a on VD CD ro sμ in I^s- en CM sμ co CM ro sμ vo oo H CM co vo r- oo sμ sμ sμ in m m in m VD vo vo I^s- I^s- c— I- t- oo CO oo co oo co

CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM t- I- I- r- L— I^s- IT- IT- I^s- I- I^s- r- I- I^s- I^s- I^s- t- r- I- I- I^s- I-

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 O O O O O O O O O O O O O O O O O O O O O O EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH t f f fcr; ( fi ^ f ι ^ ( fiai ι^ f^ f^ f^ fiai ι =ι; f f ATOM 7290 CA ALA A 542 37.066 55.364 40.230 1.00107.88 C

ATOM 7292 CB ALA A 542 35.904 55.800 39.307 1.00107.78 C

ATOM 7296 C ALA A 542 37.864 54.216 39.577 1.00108.56 C

ATOM 7297 O ALA A 542 37.628 53.033 39.876 1.00109.40 O

ATOM 7298 N VAL A 543 38.796 54.542 38.692 1.00108.53 N

ATOM 7300 CA VAL A 543 39.577 53.508 38.041 1.00108.64 C

ATOM 7302 CB VAL A 543 40.195 54.061 36.754 1.00109.00 C

ATOM 7304 CGI VAL A 543 41.305 53.152 36.228 1.00109.11 C

ATOM 7308 CG2 VAL A 543 39.092 54.231 35.693 1.00109.28 C

ATOM 7312 C VAL A 543 40.621 52.984 39.031 1.00108.59 C

ATOM 7313 O VAL A 543 40.998 51.794 39.038 1.00107.69 O

ATOM 7314 N ALA A 544 41.034 53.880 39.912 1.00108.99 N

ATOM 7316 CA ALA A 544 41.989 53.551 40.954 1.00109.92 C

ATOM 7318 CB ALA A 544 42.708 54.824 41.399 1.00109.81 C

ATOM 7322 C ALA A 544 41.342 52.846 42.161 1.00110.57 C

ATOM 7323 O ALA A 544 41.931 52.779 43.231 1.00110.09 O

ATOM 7324 N GLU A 545 40.129 52.337 41.981 1.00111.74 N

ATOM 7326 CA GLU A 545 39.370 51.695 43.056 1.00112.53 C

ATOM 7328 CB GLU A 545 38.144 52.549 43.442 1.00112.44 C

ATOM 7331 CG GLU A 545 36.842 52.200 42.714 1.00111.87 C

ATOM 7334 CD GLU A 545 35.747 53.222 42.937 1.00111.18 C

ATOM 7335 OE1 GLU A 545 35.721 53.815 44.045 1.00111.66 O

o u o a u u u o a u u u u u o a u u u u u u

cn vo H CM CM o sμ co CM ∞ sμ sμ on o vi sμ VD in I^s- CM VD vi oo cn CM sμ o o in ro CM vo vi oo o ro VD oo sμ CM o sμ o If) co ro co s m m m in VD vo r- I^s- r- VD VD in in in m sμ cn If) o rH rH rH vi H H rH rH H rH vi rH rH rH rH vi rH vi H vi rH

H H H v-i vi vi H vi H H H i vi H H H vi H vi vi vi H o O o CD o CD o o O o O o CD o O σ o o CD CD CD o o o o CD σ D o o O o O o CD o o σ o o CD CD CD o

H H H v-i H - H H H H H H vi H H vi H v- H H H vi rH

H vi H sμ ro ∞ en o rH CM VD if) in I^s- vi sμ sμ H vi sμ I^s- m H VO ir- CM co sμ VD cn vi H CM if) CM vi cn on on sμ cn sμ sμ sμ O VD sμ sμ oo i^rcn sμ CO H 00 on o CD vi VD m CM if) H sμ cn

CM CM ro rH σ on o o on on irvo I^s- CD CD σ H H o cn o CM sμ sμ sμ sμ sμ co sμ sμ ro ro on cn ro sμ sμ sμ sμ sμ sμ cn sμ sμ co oo cn o vi D 00 sμ ro irCM on co I^s- rH in in o ∞ I^s- o CM

CM vo ro rH vi H cn VD vi on oo sμ sμ cn cn H CM o CM m 00 en sμ CM ro H oo o cn r- t- o oo o CM cn on H H CM VD o VD cn ro o (D O oo en I- VD 00 oo oo co cn I^s- vo on cn O on en o σn m m sμ m sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ in sμ sμ m sμ

00 o VD cn I- oo sμ sμ CM CM m vo vo rH VD H I^s- sμ on sμ on m vi vo oo vo CM o H sμ cn vi cn vo in CM vi cn I^s- cn H ro H co D on o sμ σ o CM CM H cn vo vo sμ m vi oo on cn CM oo cn ro sμ oo on CO oo iren cn o vi H CM o CM cn CM ro sμ vo m I^s- cn ro on ro ro on on ro ro sμ sμ sμ sμ sμ sμ s sμ sμ sμ sμ sμ sμ sμ

in ιn m vo vo vo VD VD I^s- t- I^s- I^s- I^s- I- t- oo oo co oo oo oo co sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ in m in in m in in in in m m m in m m m in in m in in m ft fit fit ft fit fit fit fit fit fit fit fit ft fit fit ft fit fit fit fit ft ft

P P P ft fit fit fit fit μQ μQ ^μQ μQ μQ μQ P P P P P P P μQ μQ μQ μQ μQ μQ μQ μQ fiϋ fiϋ fiϋ fiU ^ f & PQ PQ PQ PQ PQ PQ PQ

0 0 0 f ft f f ft > > > > > > > μQ μQ μQ μQ μQ μQ μQ

CM H CM H CM

PQ f ffl fit, PQ CD CD f ffl CD P P o U O a U U U O a U U U U a U U U U u U

VD I^s- oo on vi ro t- 00 on H ro in cn cn sμ in t- D CM sμ oo CM ro ro ro ro sμ sμ sμ sμ sμ m in in in vo VD VD vo vo 1- iriroo ro cn ro ro ro ro ro on cn ro ro ro ro ro ro on on ro ro on on ro

I- I^s- I^s- r- t- I^s- r- c- I- I^s- I^s- t- I^s- r- L- I- ir- I^s- I^s- t- I^s- I^s-

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 b EH bEH bEH bEH bEH bEH EbH bEH EbH oEH oEH oEH oEH oEH oEH oEH oEH oEH oEH oE-J oH oEH ft fit fit ft fit fit fit fit fit fit fit fit fit fit fit fit fit fit fit ft ft fit ATOM 7383 0 LEU A 548 43.763 50.354 43.590 1.00115.55 O ATOM 7 7338844 N LYS A 549 42.450 48.533 43.362 1.00115.56 N ATOM 7386 CA LYS A 549 41.665 48.700 44.609 1.00115.34 C ATOM 7388 CB LYS A 549 41.400 47.325 45.281 1.00115.23 C ATOM 7391 CG LYS A 549 42.658 46.392 45.451 1.00114.71 C ATOM 7394 CD LYS A 549 42.486 44.949 44.895 1.00112.45 C ATOM 7397 CE LYS A 549 42.891 44.814 43.429 1.00111.11 C ATOM 7400 NZ LYS A 549 41.702 45.016 42.581 1.00109.06 N ATOM 7404 C LYS A 549 42.279 49.692 45.599 1.00115.43 C ATOM 7 7440055 O LYS A 549 42.859 49.278 46.594 1.00115.78 O ATOM 7 7440066 N ASP A 550 42.133 50.995 45.309 1.00115.48 N ATOM 7408 CA ASP A 550 42.718 52.101 46.103 1.00115.32 C ATOM 7410 CB ASP A 550 43.985 52.667 45.426 1.00115.13 C ATOM 7413 CG ASP A 550 45.280 52.155 46.057 1.00114.54 c ATOM 7414 OD1 ASP A 550 46.237 52.955 46.208 1.00113.39 0 ATOM 7415 OD2 ASP A 550 45.433 50.972 46.421 1.00112.42 0 ATOM 7416 C ASP A 550 41.725 53.253 46.374 1.00115.54 c ATOM 7 7441177 O ASP A 550 40.646 53.324 45.769 1.00115.42 0 ATOM 77441188 N LYS A 551 42.142 54.170 47.256 1.00115.83 N ATOM 7420 CA LYS A 551 41.252 55.139 47.907 1.00115.89 C ATOM 7422 CB LYS A 551 40.714 54.525 49.215 1.00115.59 C ATOM 7425 CG LYS A 551 41.733 53.643 50.006 1.00113.86 C

ATOM 7428 CD LYS A 551 41..055 52.665 50.970 1.00111.41 C

ATOM 7431 CE LYS A 551 40, .488 51.436 50.252 1.00110.09 C

ATOM 7434 NZ LYS A 551 39. .147 51.663 49.626 1.00107.09 N

ATOM 7438 C LYS A 551 41. .954 56.505 48.196 1.00116.75 C

ATOM 7439 0 LYS A 551 42. .887 56.580 49.010 1.00116.75 O

ATOM 7440 N LYS A 552 41. .488 57.572 47.521 1.00117.39 N

ATOM 7442 CA LYS A 552 42. .027 58.960 47.643 1.00117.39 C

ATOM 7444 CB LYS A 552 43. .416 59.064 46.959 1.00117.50 C

ATOM 7447 CG LYS A 552 44. .324 60.256 47.378 1.00117.98 C

ATOM 7450 CD LYS A 552 43. .818 61.645 46.902 1.00118.89 C

ATOM 7453 CE LYS A 552 43. .614 61.799 45.365 1.00118.83 C

ATOM 7456 NZ LYS A 552 42, .653 62.919 45.034 1.00117.84 N

ATOM 7460 C LYS A 552 41, .005 59.991 47.050 1.00116.95 C

ATOM 7461 0 LYS A 552 40. .783 60.029 45.832 1.00116.74 O

ATOM 7462 N LYS A 553 40. .391 60.811 47.914 1.00116.45 N

ATOM 7464 CA LYS A 553 39. .213 61.629 47.534 1.00116.04 C

ATOM 7466 CB LYS A 553 37. .870 60.967 47.980 1.00116.33 C

ATOM 7469 CG LYS A 553 37. .894 60.120 49.307 1.00115.81 C

ATOM 7472 CD LYS A 553 36. .944 58.891 49.297 1.00115.36 C

ATOM 7475 CE LYS A 553 37. .250 57.844 48.182 1.00114.99 C

ATOM 7478 NZ LYS A 553 38. .269 56.846 48.591 1.00114.20 N

ATOM 7482 C LYS A 553 39. .313 63.048 48.079 1.00115.31 C

ATOM 7483 O LYS A 553 38..498 63,.472 48..906 1..00114..89 0

ATOM 7484 N ASP A 554 40. .309 63. .781 47. .592 1. .00114. .63 N

ATOM 7486 CA ASP A 554 40. .631 65. .113 48. .120 1. .00114. .06 C

ATOM 7488 CB ASP A 554 41. .906 65, .083 49. .015 1. ,00114. .25 C

ATOM 7491 CG ASP A 554 41. .982 63, .877 49. .966 1. .00114. .46 C

ATOM 7492 OD1 ASP A 554 41, .257 63. .852 50. .977 1, .00114. .44 O

ATOM 7493 OD2 ASP A 554 42. .787 62. .927 49. .804 1. .00115. .81 O

ATOM 7494 C ASP A 554 40. .865 66, .125 46. .981 1. .00113. .01 C

ATOM 7495 O ASP A 554 41, .815 66, .924 47. .030 1. .00112. .70 O

ATOM 7496 N LEU A 555 40. .024 66. .098 45, .955 1. ,00111, .82 N

ATOM 7498 CA LEU A 555 40. .272 66 . .969 44. .807 1. .00111. .02 C

ATOM 7500 CB LEU A 555 41. .150 66. .244 43. .774 1. ,00111. .12 C

ATOM 7503 CG LEU A 555 42. .365 66, .982 43. .171 1. .00111. .63 C

ATOM 7505 CD1 LEU A 555 42. .974 68. .057 44. .104 1. .00110. .98 C

ATOM 7509 CD2 LEU A 555 43. .436 65. .951 42. .717 1. ,00112. .19 C

ATOM 7513 C LEU A 555 39. .012 67. .540 44. .143 1. .00109, .95 c

ATOM 7514 O LEU A 555 37. .902 67, .001 44. .279 1. .00110, .06 O

ATOM 7515 N SER A 556 39. .226 68, .653 43. .442 1. .00108, .08 N

ATOM 7517 CA SER A 556 38. .185 69 . .374 42. .719 1. .00106. .68 C

ATOM 7519 CB SER A 556 37. .222 70, .080 43. .695 1. ,00106. ,67 C

ATOM 7522 OG SER A 556 37. .779 70, .240 44. .993 1. .00106. .13 O

ATOM 7524 C SER A 556 38. .900 70, .382 41. .832 1. .00105. .37 C

o a u u u u u u o u u u o a u u u u a u a a

sμ r- VD if) sμ CM sμ 00 I^s- CD VD o cn cn oo in CO VO cn o r- VD sμ o o cn CM r- CM CM sμ sμ ro o CD on ro σn vo sμ CM ro rH σ m sμ ro CM CM vi σ cn r- vi H CM vi o O σ vi CM m vo vo sμ σ o o σ o o o cn cn o o O o o O σ CD o o o o o rH vi H H H H H rH H rH H H vi vi i H vi H H H o CD o σ o o σ o o σ o O σ O CD O CD o O O o o σ CD o σ o o σ o o o o O o O CD σ CD o σ O o o

H i H vi vi H H H vi H H vi H H rH H H vi vi i rH vi cn sμ VD CD sμ m VD H VD on sμ CD ID in sμ s VO sμ cn vi O oo oo I^s- on if) CM vo r- ro ro vo vo VO H vo m I^s- VD sμ t- sμ 00 sμ

CM m vo i vo H VD VD rH σ in H cn oo r- CM H I^s- I^s- o CM σ

CM o en ∞ I^s- irVD VD VO r- I^s- O cn o vi CM H H o H CM σ sμ sμ ro ro ro on ro ro ro oo ro sμ ro sμ sμ sμ sμ sμ sμ sμ sμ sμ i o ro vi I^s- cn co oo on 00 I^s- ^r- sμ H in t- O vi H on o H

I^s- VD oo t- t- ro I- in ro CM oo co VD VD ∞ in H cn VO sμ o I^s- on m in ro VD VD oo H ro CM on on on o H σ i cn σ sμ o CM o o H vi H o σ CM CM cn CM CM ro cn sμ sμ sμ sμ sμ ro co ro irt- trt- I^s- i- I^s- r- irI^s- I^s- I— I^s- r- I^s- I^s- I^s- I^s- I^s- I^s- I^s- iron ro io sμ vo CM cn vo on VD σn I^s- σ 00 i cn H co CM cn CM on in oo rH I^s- VD sμ sμ VD o m m cn on D I^s- 00 ro cn r- oo oo σ oo sμ vi oo sμ vo cn co vo VD on vo ro in CM co t- n CM o I^s- CM cn co CD oo I^s- VD in sμ cn in vo oo on irI^s- in sμ cn CM vi o σ ro cn cn cn ro co cn m cn ro cn cn ro on ro cn n cn ro cn cn ro

vo r- r- t- t- r- I^s- r- I^s- I^s- r- r- I^s- ∞ oo oo ∞ oo oo oo oo ∞ in m m in m m in m in m ιn m ιn in m in m m in in m m in in m in m in m m in in in m in in m if) in m in in m in ft fit fit fit fit fit fit fit fit fit fit fit fit fit ft fit fit fit fit fit fit fit pi pti Pi Pi Pi Pi pi Pti Pi Pi Pi pi Pi ø ø ø ø ø ø ø ø ø

H H H H 1 txH txH txH ^■xH txH txH !χH txH p p^ - 0-: C_{- -}i p_; 0-. _: co EH EH EH EEHn EEHH H E EHπ HE^H fcEHH HE^H EEH^H f-ϋ f-ϋ ril fiϋ fi fiϋ fiϋ fiϋ fiϋ vi H CM CM H CM riJ ffl ø P PQ ISl lDl Liq p ft, ffl 0 o a u u u u u u o u u u o a u u u Pu PaQ IuS] Pai Xa in D oo o fO sμ vo oo on rH 00 m vo I^s- on rH sμ tt-- o CM ro ID

CM CM CM ro ro ro ro ro ro sμ sμ sμ sμ sμ sμ in m mm VD VO VD VD m m in in in m in in m in m in in m in m m iinn in in m in

I- I^s- I^s- I^s- I^s- I^s- t- i^rI^s- I^s- I^s- I^s- I^s- irI^s- tr- 1- rr-- IT- t- r- I-

o 2 2 2 2 o2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2

EH oEH oEH oEH EH oEH oEH oEH bEH bEH bEH bEH bEH bEH bEH bEH bEH oEH oEH oEH oH oEH fi fi fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ fi fiϋ fi fiϋ fiϋ fi fi fi fi u o a u u u o o u o a u u u u u u o a u u u

in σn CD oo on CM I^s- CM on VD i ro sμ i in cn oo cn sμ VD o VD trH CO 00 ∞ r- ro oo CM in in CM cn sμ H cn in if) m VD cn cn io m m VD vo ιn ro cn o I- I^s- oo cn vo m cn cn cn ro ro cn on on on en on on en σn on cn oo oo oo co co ∞ oo oo co 00 oo o o o o o o o o o o o σ o σ o σ o o o o o o o o o o o o o o o σ o σ o σ o o o o o o o o vi H H H H r-i vi H H H vi H H vi H H vi vi H H vi vi

CM vo cn CM oo sμ r- VO sμ oo CM o r- CM I^s- H CM oo CM ro cn co cn VD irCM vo D cn m oo VD ro ro H iT- vo CM in CM on o sμ m oo I^s- on H VD sμ o sμ oo trr- ro on cn VD on ro on r- I^s- cn H

CM CM ro in m vo VD t- sμ io 00 ro vi H O H ΓO cn CM CM vi rH sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ s}< o m CM oo o sμ on in o vi m ro CM rH VD m in CM o in CM O ro oo cn I^s- cn in ro oo VD cn en H I^s- I^s- CM sμ D 00 n oo if) co v n oo r- cn on sμ H CM on ∞ ro I^s- CM m sμ r- CM ) cn sμ CM sμ cn sμ sμ cn CM ro CM m in sμ m sμ 00 CM in o I^s- r- oo D oo

I^s- I^s- I^s- trI^s- I^s- r- t- t- I^s- I^s- I^s- r- r- r- I^s- I- I^s- I^s- I^s-

H VD r io I^s- o CM vo o CM on CM o CM H VD ∞ o on I^s- CM o H oo VD H VD CM H VD r- VD vo CM I^s- CM sμ I^s- 00 rH on sμ o o ro CM H H rH on 00 on in H i m oo H t- σ vo VD I^s- cn I^s- ro oo CD co on on I^s- VO t- o H vi CM CM CO CM sμ CM cn H o o cn cn ro ro co ro cn n sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ

oo ∞ cn cn cn cn on cn cn cn o o o σ o o o o H vi H H m in in in m in in in m in vo VD vo VD VD 10 vo vo vo 10 VD vo m in m m m in m m in in m m m in in in in m in in m m ft ft fit fit fit fit fit ft fit fit fit ft ft fit fit fit fit fit fit fit fit fit

CD ϋ ft ft ft ft ft ft ft ft PQ PQ PQ PQ PQ PQ PQ PQ O O O O Pi P-i tO CO CO CO CO tO tO CO P P μQ μQ μQ P P P Pi Pi Pi P-; rtl riJ fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ H H H H H H H H ft ft ft ft

on σ H co m CO cn o vi CM cn in I- on CM VD o H CM cn m ∞ vo t- I^s- l— I^s- I- r- oo oo oo oo ro ∞ ∞ en on o O O o o o m ιn m m in in m in in in m m m m m m vo VD VO o vo vo

I^s- t- r- t- I^s- t- I^s- I- I^s- I^s- r- I^s- I^s- I^s- I- r- I- I^s- I^s- I- r- I^s-

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 O O O O O O O O O O O O O O O O O O O O O O EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH ft ft fit fit fit fit fit fit fit fit fit fit fit fit ft fit ft fit fit fit fit fit u u o a u u u o a u u u u u u o u u u o a u

m vi o oo ro I^s- I^s- H in o oo sμ in VD on m CM CM in on I^s- o sμ en CM VD vo I^s- ro sμ o cn H CM ro o sμ I^s- I^s- VD sμ vo sμ rH ro CM cn H o o cn on ∞ VD I^s- (D CM m sμ sμ CM O sμ CM o oo co co ∞ co oo I^s- r- r- I^s- I- r- CO ∞ oo CO ∞ co I- I^s- r- IT- o o o o o o o o o o o o o o o o o o o o o CD o o o o o o o o o o o o o o o o o o o o o o vi rH rH vi H vi H H H H H H rH v-i H vi rH H H H vi rH cn ir- sμ cn vi if) CM vi oo VD sμ H in co vi cn I^s- VD I^s- O o CM o co I- if) vo sμ CM CM sμ I- I^s- sμ I- r- CM ir- o CM sμ m CM en o o H vi sμ CM CM o on sμ CM cn VD CM H vo sμ CO oo in m I^s-

CM sμ sμ in vo I^s- I^s- oo vo I- I^s- I^s- oo CD cn on ∞ I^s- vo I- in sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ ji m vo CM ID VD ro VD cn cn irm VD oo in CM 1- vo I- rH VD cn o cn in cn oo I^s- vo ro o vo on o oo oo cn cn ro oo oo in on cn m o vo I^s- cn m in on VD vo o in m o vi ιr- en CM vi vo cn r- m

I^s- cn o co on on co on r- t- m sμ in sμ CM H CM cn I^s- I^s- r- co I^s- r- oo ir- I^s- I^s- i^rI^s- r- t- I^s- I^s- I^s- I- t- I^s- I^s- ir- I^s- I^s- I^s- ircn ιn I^s- co VD sμ on CM H ro H sμ sμ I- CM m o co I^s- rH co on o sμ H ro ro ro o Lf) CM I- oo vo vo CM cn ir- co o H CM vo oo

VD o in vo CO m o VD ro m vo VD in VD I^s- co ∞ CO oo I^s- oo oo o CM CM vi vi o ro cn ro sμ sμ ro CM H vi o CM ro m VD in VD sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ

H rH H CM CM CM CM CM ro ro co ro ro n ro ro on ro ro ro sμ sμ

VD VD vo VD vo VD VO VD vo vo VD vo D VD VD vo VD vo VD vo vo VD in m m m m m in in in in in in m in in in m in m m m m fit fit ft ft fit fit fit fit fit fit << ft ft fiU fiϋ f fiϋ fit ft fiϋ fiϋ

O O O ft fit fit fit fit Pi Pi Pi Pi Pi Pi Pi Pi Pi Pi Pi Pi Pi Pi

Pi Pi Pi μQ μQ μQ μQ μQ tx tx tx tx tx tx tx tx tx tx tx tx ^■x tx ft ft ft fit fit ft ft fit fcn fcH fcH fcH H fcH fcH EH EH H EH EH

O fit, Q fit ffl CD P PQ N Pi PQ P fit

U U O a U U U O a υ U υ υ υ U O U u U b a U

H sμ in vo oo o sμ in VD oo o ro sμ vo co on H ro in vo r- CD

H rH vi H vi CM CM CM CM CM cn ΓO ro ro ro ro sμ sμ sμ sμ sμ sμ

VD vo vo o vo VO VD VD VD VD D vo vo vo VD vo VD vo VD vo vo vo

I^s- I- r- I^s- r- I- I- I^s- r- ir- r- I i^rI^s- I- irI- t- I^s- I^s- r- I^s-

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 O O O O O O O O O O O O O O O O O O O O O O EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH ft fit fit fit fit fit fit fit fit fit fit fit fit fit fit fit fit ft fit fit fit fit u u u u u o u u u o a u u u o a u u u o a u

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2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 O O O O O O O O O O O O O O O O O O O O O O H EH EH EH EH H EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH ft fit fit fit fit fit fit fit fit fit fit fit fit fit fit ft ft fit fit fit fit fit ATOM 9507 CA VAL A 743 32,.100 33..380 -15..780 1..00120..00 c

ATOM 9509 CB VAL A 743 31. .349 34. .588 -15. .092 1. .00119. .99 c

ATOM 9511 CGI VAL A 743 32. .075 34. .985 -13. .812 1. .00120. .37 c

ATOM 9515 CG2 VAL A 743 31. .133 35. .795 -16. .045 1. .00120. .08 c

ATOM 9519 C VAL A 743 31. .638 32. .155 -14. .993 1. ,00119. .99 c

ATOM 9520 0 VAL A 743 32. .460 31. .345 -14. .542 1. .00119. .82 0

ATOM 9521 N ALA A 744 30. .315 32. .081 -14. .798 1. .00119. .83 N

ATOM 9523 CA ALA A 744 29. .661 31, .009 -14, .072 1. .00119. .78 C

ATOM 9525 CB ALA A 744 28. .470 31. .564 -13. .241 1. .00119. .49 C

ATOM 9529 C ALA A 744 29. .201 29. .900 -15. .056 1. .00119. .83 C

ATOM 9530 0 ALA A 744 29. .311 28. .706 -14. .715 1. .00120. .22 0

ATOM 9531 N HIS A 745 28. .727 30. .284 -16. .262 1. .00119. .24 N

ATOM 9533 CA HIS A 745 28. .096 29. .348 -17. .241 1. .00118. .60 C

ATOM 9535 CB HIS A 745 27, .483 30. .107 -18. .463 1. .00118. .61 C

ATOM 9538 CG HIS A 745 26 .070 30, .609 -18, .247 1. .00119. .45 C

ATOM 9539 ND1 HIS A 745 25 .452 31, .496 -19, .110 1. .00119. .14 N

ATOM 9541 CE1 HIS A 745 24, .234 31. .764 -18. .670 1. .00119. ,23 C

ATOM 9543 NE2 HIS A 745 24, .032 31. .084 -17. .553 1. .00119. .61 N

ATOM 9545 CD2 HIS A 745 25, .161 30. .351 -17. .268 1. .00120. .04 C

ATOM 9547 C HIS A 745 29, .004 28. .193 -17. .718 1. .00117. .70 C

ATOM 9548 0 HIS A 745 28, .493 27. .137 -18. .080 1. .00117. .16 O

ATOM 9549 N LEU A 746 30 .325 28, .393 -17, .682 1. .00117. .07 N

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2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 O O O O O O O O O O O O O O O O O O O O O O EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH ft ft fit fit fit fit fit ft fit fit fit fit fit fit fit fit ft ft ft ft fit ft a u u u o a u u u u a u a a u o a u u u u u

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VD ir- VD ir- t- VD L o cn vi CM co H sμ cn cn vi on o vi o I- oo co 00 co 00 00 oo co ∞ on σn cn σn on oo oo oo I- co co oo t-- o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o σ o o o o o o o o o

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H vi H vi vi vi H vi rH rH v-i r-t H H H H vi H H H H v-i

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 O O O O O O O O O O O O O O O O O O O O O O EH EH EH EH EH EH EH H EH EH EH EH EH EH EH EH H EH EH EH EH EH ft ft fit fit fit fit rt fit fit fit fit fit fit fit fit fit ft fit fit fit fit ft o a u u u u u u o a u u u u u o a u u u u o

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00 r- VD m LO sμ sμ vo r- LO sμ cn 00 sμ sμ sμ LO in in t- ir- t-^r 1 1 1 1 1 1 1 1 1 1 1 I 1 1 1 1 1 1 1 1 1

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O o o o D o o o o o o o o o o o o o σ o o O

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H CM H fiϋ ffl CD P P fiϋ ffl CD P fiϋ ffl 0 P PQ o a U U U U U U O a U U U U u O a u u U U o v-i CM sμ VD on vi m on o H CM sμ I^s- o ro sμ LO t- σn CM LO VD cn cn ro ro cn sμ sμ sμ m LO LO io lo VD vo vo VD VD ID I- tr- I- vo VD vo vo VD VD VD VD vo VD VO vo vo VD VD VD VD VO vo VD VD vo cn ro ro ro on cn ro ro ro ro CO ro ro ro ro ro ro ro on on cn ro vi H H H H vi H vi H H H vi H vi vi H H H H H H H

2b b2 b2 2b 2b b2 o2 o2 2o 2o 2o o2 2b 2b 2b 2b 2b o2 2o 2o 2o 2o

EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH ft ft fit ft fit fit fit fit fit fit ft fit fit ft fit fit fit fit fit fit fit fit ATOM 13677 OE2 GLU A1005 8..971 37..513 43..652 1..00 75..46 0

ATOM 13678 C GLU A1005 3, .787 39. .289 41, .243 1. .00 68. .88 c

ATOM 13679 O GLU A1005 3. .606 39. .995 42. .236 1. .00 68. .88 0

ATOM 13680 N GLY A1006 2. .804 38. .681 40. .578 1. ,00 66. .86 N

ATOM 13682 CA GLY A1006 1. .481 38. .559 41, .172 1. .00 66, .23 C

ATOM 13685 C GLY A1006 0. .700 39. .863 41, .245 1. .00 64. .99 C

ATOM 13686 0 GLY A1006 0. .307 39. .986 41, .974 1. .00 64, .16 O

ATOM 13687 N LEU A1007 1. .212 40. .831 40, .479 1. .00 63, . 66 N

ATOM 13689 CA LEU A1007 0. .593 42. .141 40. .313 1. ,00 61. .90 C

ATOM 13691 CB LEU A1007 1. .108 42. .815 39, .046 1. .00 61. .68 C θJ

)—- ATOM 13694 CG LEU A1007 0. .564 42. .140 37, .775 1. .00 59. .69 C

ATOM 13696 GDI LEU A1007 1. .048 42. .762 36. .455 1. .00 57. .45 C

ATOM 13700 CD2 LEU A1007 0. .952 42. .157 37. .860 1. .00 58, .64 c

ATOM 13704 C LEU A1007 0. .739 43. .080 41. .463 1. .00 60. .23 c

ATOM 13705 O LEU A1007 0. .025 44, .000 41, .508 1. .00 60, .14 O

ATOM 13706 N GLN A1008 1, .654 42, .810 42, .389 1. .00 58. .70 N

ATOM 13708 CA GLN A1008 1. .834 43. .562 43. .643 1. ,00 58. .39 C

ATOM 13710 CB GLN A1008 2. .374 42. .639 44. .751 1. 00 60. .04 C

ATOM 13713 CG GLN A1008 3. .728 42, .962 45. .376 1. ,00 64. .86 C

ATOM 13716 CD GLN A1008 4. .555 41. .661 45. .640 1. ,00 73. .84 C

ATOM 13717 OE1 GLN A1008 4. .215 40. .869 46. .546 1. ,00 79. .05 O

ATOM 13718 NE2 GLN A1008 5, .613 41. .434 44. .836 1. ,00 74. .59 N

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ATOM 13900 CZ ARG A1020 8. .616 55. .398 51, .262 1. .00 86, .95 C

ATOM 13901 NH1 ARG A1020 8. .309 54. .986 50. .017 1. .00 87. .49 N

ATOM 13904 NH2 ARG A1020 9. .735 54. .951 51, .845 1. .00 85, .66 N

ATOM 13907 C ARG A1020 4. .715 60, .806 50, .123 1. .00 64, .44 C

ATOM 13908 O ARG A1020 5. .461 61. .502 50, .807 1. .00 63, .43 O

ATOM 13909 N ILE A1021 3, .939 61, .296 49, .162 1. .00 64, .78 N

ATOM 13911 CA ILE A1021 3, .900 62. .687 48, .765 1. .00 65. .83 C

ATOM 13913 CB ILE A1021 3. .222 62. .788 47. .351 1. .00 65. .53 C

ATOM 13915 CGI ILE A1021 3. .999 62. .073 46. .279 1. .00 64. .95 C

1-0 -4 ATOM 13918 CD1 ILE A1021 3. .140 61. .974 45. .018 1. .00 63. .31 C

ATOM 13922 CG2 ILE A1021 3. .032 64. .249 46. .873 1. .00 65. .37 C

ATOM 13926 C ILE A1021 3. .076 63. .565 49. .720 1. .00 67. .43 C

ATOM 13927 O ILE A1021 3. .530 64. ,600 50. .171 1. .00 66. .80 O

ATOM 13928 N THR A1022 1. .821 63. .176 49. .949 1. .00 69. .95 N

ATOM 13930 CA THR A1022 0, .864 64, .017 50, .604 1. .00 72. .27 C

ATOM 13932 CB THR A1022 -0. .601 63. .630 50. .298 1. ,00 71. .62 C

ATOM 13934 OG1 THR A1022 -0. .790 62. .222 50. .394 1. .00 70. .87 O

ATOM 13936 CG2 THR A1022 -0. .957 63. .924 48. .951 1. ,00 72. .16 C

ATOM 13940 C THR A1022 0. .985 63. .968 52. .070 1. .00 75. .93 C

ATOM 13941 O THR A1022 0. .103 64. .496 52. .733 1. ,00 77. .38 O

ATOM 13942 N ASP A1023 2. .011 63. ,368 52. .649 1. .00 80. .01 N

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ATOM 15317 CH2 TRP A1196 -4. .180 57. .393 48, .106 1. .00 67, .03 c

ATOM 15319 CZ2 TRP A1196 -5. .143 58. .317 48. .199 1. .00 65. .64 c

ATOM 15321 C TRP A1196 -5. .421 60. .667 41. .154 1. .00 59. .03 c

ATOM 15322 0 TRP All96 -4. .189 60. .672 41. .027 1. .00 58. .55 0

ATOM 15323 N HIS A1197 -6. .250 60. .637 40, .107 1. .00 59. .19 N

ATOM 15325 CA HIS A1197 -5. .765 60. .643 38, .703 1. .00 59. .56 G

ATOM 15327 CB HIS A1197 -6. .888 60. .238 37, .708 1. .00 58, .43 C

ATOM 15330 CG HIS A1197 -7. .276 58, .803 37, .766 1. .00 53, .66 C

ATOM 15331 ND1 HIS A1197 -6, .605 57. .867 38, .545 1. ,00 49, .63 N

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ATOM 15335 NE2 HIS A1197 -8, .214 56, .829 37 .570 1. .00 50, .51 N

ATOM 15337 CD2 HIS A1197 -8. .299 58, .149 37, .175 1. .00 49, .73 C

ATOM 15339 C HIS A1197 -5. .252 62, .034 38, .314 1. ,00 60, .86 C

ATOM 15340 0 HIS A1197 -4. .086 62. .197 37. .843 1. .00 62. .76 O

ATOM 15341 OXT HIS A1197 -6, .124 62, .910 38. .493 1. .00 61. .16 O

ATOM 15342 N THR B 59 58. .236 -29, .201 -11, .478 1. .00 66. .12 N

ATOM 15344 CA THR B 59 59, .426 -29, .069 -10, .536 1. .00 66. .43 C

ATOM 15346 CB THR B 59 60. .208 -27, .735 -10, .791 1. .00 66. .42 C

ATOM 15348 OG1 THR B 59 60. .220 -27, .425 -12, .190 1. .00 66. .16 O

ATOM 15350 CG2 THR B 59 61 .744 -27, .849 -10, .416 1. .00 65. .35 C

ATOM 15354 C THR B 59 59 .073 -29, .185 -9, .003 1. .00 66. .30 C

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ATOM 18123 N ARG B 241 74, .072 -21. .139 16 .332 1. .00 99 .68 N

ATOM 18125 CA ARG B 241 74. .821 -22. .206 15, .692 1. .00100, .43 C

ATOM 18127 CB ARG B 241 73. .858 -23. .371 15, .412 1. .00100. .60 C

ATOM 18130 CG ARG B 241 73, .449 -24. .142 16 .687 1. .00102 .62 C

ATOM 18133 CD ARG B 241 74. .565 -25. .079 17, .313 1. .00105, .74 C

ATOM 18136 NE ARG B 241 74, .127 -25. .730 18 .577 1. .00107 .69 N

ATOM 18138 CZ ARG B 241 74. .921 -26. .114 19, .616 1. ,00107. .81 C

ATOM 18139 NH1 ARG B 241 76. .246 -25. .946 19. .598 1. ,00106. .93 N

ATOM 18142 NH2 ARG B 241 74. .369 -26. .665 20, .705 1. .00106. .86 N

ATOM 18145 C ^' ARG B 241 75, .637 -21. .708 14. .426 1. .00100. .28 C

ATOM 18146 0 ARG B 241 76, .009 -20. .525 14, .380 1. .00100, .31 O

ATOM 18147 N GLY B 242 75, .980 -22. .601 13, .471 1. .00 99. .88 N

ATOM 18149 CA GLY B 242 76, .502 -22. .235 12, .150 1. .00 99, .12 C

ATOM 18152 C GLY B 242 77, .842 -21. .547 12, .204 1. .00 98, .82 C

ATOM 18153 0 GLY B 242 78. .445 -21. .544 13. .255 1. ,00 98. .92 O

ATOM 18154 N ILE B 243 78, .313 -21. .000 11. .082 1. .00 98. .44 N

ATOM 18156 CA ILE B 243 79. .499 -20. .122 11. .039 1. ,00 98. .44 C

ATOM 18158 CB ILE B 243 80, .349 -20. .324 9, .701 1. .00 99. .38 C

ATOM 18160 CGI ILE B 243 81, .300 -21. .554 9, .744 1. .00100. .12 c

ATOM 18163 CD1 ILE B 243 81. .538 -22. .272 8. .329 1. .00100. .08 c

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ATOM 22315 0 GLN B 510 52.692 -28.842 26.642 1. .00 96. .57 O

ATOM 22316 N LYS B 511 50.540 -29.402 26.968 1. .00 93. .52 N

ATOM 22318 CA LYS B 511 50.176 -29.304 25.549 1. .00 91. .61 C

ATOM 22320 CB LYS B 511 48.849 -30.016 25.256 1. .00 91. .42 C

ATOM 22323 CG LYS B 511 48.927 -31.523 25.300 1. .00 91. .13 C

ATOM 22326 CD LYS B 511 47.629 -32.151 25.879 1. .00 90. .90 C

ATOM 22329 CE LYS B 511 47.818 -33.645 26.201 1. .00 89. .31 C

ATOM 22332 NZ LYS B 511 49.165 -34.111 25.736 1. .00 87. .10 N

Ui -J o ATOM 22336 C LYS B 511 50.019 -27.835 25.119 1. .00 90. .54 C

ATOM 22337 0 LYS B 511 49.620 -26.955 25.937 1. .00 90. .21 O

ATOM 22338 N ARG B 512 50.327 -27.585 23.834 1. .00 88. .48 N

ATOM 22340 CA ARG B 512 49.873 -26.382 23.096 1. .00 86. .62 C

ATOM 22342 CB ARG B 512 50.826 -26.081 21.914 1. .00 86. .70 C

ATOM 22345 CG ARG B 512 52.224 -25.564 22.267 1. .00 87. .71 C

ATOM 22348 CD ARG B 512 53.386 -26.264 21.526 1. .00 89. .00 C

ATOM 22351 NE ARG B 512 53.179 -26.372 20.082 1. .00 90. .27 N

ATOM 22353 CZ ARG B 512 53.447 -27.449 19.333 1. .00 91. .27 C

ATOM 22354 NH1 ARG B 512 53.948 -28.563 19.877 1. .00 89. ,77 N

ATOM 22357 NH2 ARG B 512 53.205 -27.404 18.016 1. .00 92. .51 N

ATOM 22360 C ARG B 512 48.378 -26.528 22.564 1. .00 84. .19 C

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CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM

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CM on LO VD on o I- VD sμ ro s on t- en CM oo o H VD σn in CM

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1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 t- CM CM LO VD H on LO VD co LO oo o co r- on sμ on to vo rH H

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o o o o a o o o o a o o o o o o a o o o o o

C08910/l7002SIl/X3[ 9176801/17002 OΛV ATOM 30594 NE ARG BIO89 15.316 -46.783 54.794 1.00 91.06 N

ATOM 30596 CZ ARG B1089 14.016 -46.585 55.052 1.00 94.11 C

ATOM 30597 NH1 ARG B1089 13.229 -45.952 54.164 1.00 95.54 N

ATOM 30600 NH2 ARG B1089 13.490 -47.023 56.204 1.00 93.47 N

ATOM 30603 C ARG B1089 20.684 -44.557 52.702 1.00 83.49 C

ATOM 30604 O ARG B1089 21.426 -45.186 51.944 1.00 83.54 O

ATOM 30605 N LEU B1090 20.672 -43.230 52.767 1.00 81.93 N

ATOM 30607 CA LEU B1090 21.579 -42.429 51.973 1.00 81.44 C

ATOM 30609 CB LEU B1090 21.292 -40.949 52.140 1.00 81.96 C

ATOM 30612 CG LEU B1090 19.951 -40.525 51.548 1.00 83.88 C

-J

LΛ ATOM 30614 CD1 LEU B1090 19.576 -39.125 52.020 1.00 84.40 C

ATOM 30618 CD2 LEU B1090 19.985 -40.613 49.994 1.00 84.77 C

ATOM 30622 C LEU B1090 23.017 -42.673 52.316 1.00 80.51 C

ATOM 30623 O LEU B1090 23.827 -42.792 51.419 1.00 81.29 O ATOM 30624 N ALA B1091 23.371 -42.746 53.597 1.00 79.41 N ATOM 30626 CA ALA B1091 24.793 -42.897 53.940 1.00 78.42 C

ATOM 30628 CB ALA B1091 24.999 -42.742 55.420 1.00 78.06 C

ATOM 30632 C ALA B1091 25.306 -44.249 53.455 1.00 77.45 C

ATOM 30633 O ALA B1091 26.487 -44.473 53.290 1.00 76.08 O

ATOM 30634 N ALA B1092 24.367 -45.155 53.247 1.00 77.58 N

ATOM 30636 CA ALA B1092 24.664 -46.528 52.893 1.00 77.41 C

ATOM 30638 CB ALA B1092 23.476 -47.420 53.232 1.00 77.60 C

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VD vi CM sμ oo CD VD o CM I- v-i VD en i^rLO oo VO ^■sμ sμ H ro C- I- CM sμ o CM LO I- LO v-i sμ CM v-i on o sμ on on O LO O sμ sμ CM sμ VD LO cm ir- σn sμ sμ vi σn v-i VD ir∞ troo 00 vo CM sμ r-

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LO VD oo r- vo ro VD VD I- co ∞ cn cn o t- r- vo LO sμ sμ in CM sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ m sμ sμ sμ sμ sμ sμ sμ sμ

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 t- o CM in CM CM o on cn H on VD vi CD H on I- co oo CM CM VD o VD sμ t^r- LO I- o σn H sμ CM r- LO O I- oo CM LO 00 o VO LO sμ o o CM sμ o oo 00 oo σn VO ∞ sμ VO CM vo CM sμ vi oo sμ I- cm on o vi vi CM on o oo oo t^r- ID in I- on oo o O cn oo oo co

CM CM ro cn cn ro CM ro CM CM CM CM CM CM CM CM ro ro CM CM CM CM

LO in in LO in in LO IO VD VD vo vo VD VO vo VO irirI- r- ircm σn cm cm σn σn σn σn on on on on on on on en o on on σn σn on o o o o o o o o o o o o o o o o o o o o o o vi v-i rH rH vi rH H vi H vi v-i rH ri vi H vi H vi H H vi H ffl ffl ffl PQ ffl ffl ffl m ffl ffl ffl ffl ffl ffl ffl ffl ffl ffl ffl ffl Q ffl p p p p p p p p p p p p p p p p ffl ffl ffl ffl ffl ffl j j μ-q j j I-q W H H H H H pq w w c W W CΛ ø ø ø ø ø ø ø ø p p p p p p p p fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ

H CM H CM H CM ft ffl 0 P PQ PQ ft ffl 0 P P fit ffl 0 P P u U U U U U U U a U U U U U U U a U U U U U cm vi sμ r- oo cm o vi CM sμ vo cm H m cm o vi en in oo em o co on cm σn cn cm o CD O o o o vi vi H CM CM CM CM CM CM ro vo vo vo 10 10 0 r- r- r- r- r- r- tr- t- t- r- I- r- I- r- tr- r- o o o o o o o o o o o o o o o o O o o o o o ro ro ro cn ro ro ro ro cn ro ro ro ro cn ro ro ro ro cn ro ro ro

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 U U O O O O O O O O O O O O O O O O O O O O EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH ft fit ft fit ft fit fit fit ft ft fit fit fit ft fit ft fit fit ft fit fit fit u u a u u u u u a u u a u u u u u u u a u u

o r- cn sμ vo I- oo I- en O on oo oo sμ ro v-i sμ o oo O ro o sμ vo cn cn sμ VD vo ro cn cn sμ ro ro ro ro o -i rH on I- r- I- t- r- t- t- t- t- I- t- r- t- r- t- D vo lO o o o o o o o O o o o o o o o o o O o O o o o o o o o O o o o o o o o o o O o O v-i H i vi v- vi H ri -i vi v-i H vi vi H H vi r-i H H H v-i o r-i m o o vi o vi cn on on on I- LO o tr- on oo m L0 vi en vi vi cm o on ID C- 00 rH o oo co H on en on on ro VD 00 rH ro vo in sμ in m VD trcn o cm o on O O O on em sμ sμ vo vo VD io vo in vo VD ID sμ vi o O o sμ co r- m vi H oo rH sμ o oo rH I- m rH ro ro O H ro I- vo ro sμ v sμ sμ H oo H cn on on H on on oo on sμ sμ 00 sμ ircn on cn sμ to H o H o o sμ sμ cn 00 ro ro on ro on ro ro cn ro cn cn on ro cn ro ro ro ro

1 1 1 ro vi cm tr- I- ro ro o ro vi r- oo o on oo to oo rH sμ vi H sμ ∞ cn trt- cm sμ ro l-^r cn tr- r- vo I- en cn t- on I- cn io sμ sμ o oo O sμ cm rH oo

H cn 00 on oo r- cm oo o o cm co cm t- o o sμ cn ro vi vi H i i H r-i vi H vi H vi vi

tr- I- vo vo vo VD VD VD vo o VD rr- r- I- r- t- I- r- r- oo oo oo on σn ro cn 00 cn on on on on on on ro co ro ro oo ro ro on on on o o H vi rH H vi vi H vi rH r-i rH H vi H H H vi rH rH vi

H H H vi H vi vi vi H vi -i rH H H vi H rH H vi vi H vi ffl ffl ffl ffl ffl ffl ffl ffl ffl ffl ffl ffl ffl ffl ffl ffl ffl ffl ffl ffl ffl PQ ffl ft ω ω w c ω ω c oo Do p p p p p p p p p p p ϋ- ϋ3 H H H H >H H H H !» ρq ρq pQ pq ρq pQ PQ PQ PQ PQ pq fitJ fiC P P P P P P P P P P P P P P P P P P P P vi CM f ffl r P Q rsi f ffl to Ω P ft ffl u U a U U u U U a U U a U U U U U U υ a υ U vi CM on m r- o en VD cm ro sμ m σn CM oo CM ro sμ VD oo cn cn ro ro ro sμ sμ sμ sμ LO m LO lO to VD VD VD I- t- r- r- I- tr- tr- r- I- I- t- r- r- r- t- I- t- I- t- t- r- r- I- r- t- t- I- o o o o o o o o o o o o o o o o o o o o o o ro ro en ro ro ro en ro ro ro ro ro cn ro cn cn ro oo ro ro cn ro

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 U U U U O O O O O O O O O O O O O O O O O O IH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH ft fit ft fit fit fit ft fit fit fit fit ft fit fit fit fit fit fit fit fit fit ft ATOM 30781 CG LEU B1138 25..239 -31,.126 60,.468 1..00 69,.66 c

ATOM 30783 GDI LEU B1138 24. .909 -32, .389 61, .299 1. .00 74, .39 c

ATOM 30787 CD2 LEU B1138 26. .652 -31, .252 59, .920 1. .00 71, .45 c

ATOM 30791 C LEU B1138 22. .071 -30. .069 58, .446 1. .00 66 . .10 c

ATOM 30792 O LEU B1138 21. .069 -30. .637 57, .941 1. .00 64. .42 0

ATOM 30793 N VAL B1139 22. .582 -28. .922 57, .976 1. .00 64, .82 N

ATOM 30795 CA VAL B1139 21. .848 -28. .131 56, .970 1. .00 64. .36 C

ATOM 30797 CB VAL B1139 22. .429 -26. .716 56. .778 1. .00 64. .77 C

ATOM 30799 CGI VAL B1139 21. .545 -25. .920 55. .798 1. .00 64. .62 C

ATOM 30803 CG2 VAL B1139 22, .542 -25, .973 58, .142 1. .00 64. .98 C

ATOM 30807 C VAL B1139 21, .819 -28, .841 55, .632 1. .00 62. .71 c

ATOM 30808 O VAL B1139 20. .833 -28. .855 54. .896 1. ,00 61. .51 o

ATOM 30809 N SER B1140 22. .938 -29. .478 55. .397 1. .00 61. .64 N

ATOM 30811 CA SER B1140 23. .186 -30. .275 54. .237 1. .00 60. .62 C

ATOM 30813 CB SER B1140 24. .671 -30. .614 54. .225 1. .00 60. .69 C

ATOM 30816 OG SER B1140 24. .830 -31. .838 54. .887 1. .00 61. .55 O

ATOM 30818 C SER B1140 22, .429 -31. .593 54. .197 1. .00 59. .60 C

ATOM 30819 O SER B1140 22, .203 -32. .134 53, .120 1. .00 59. .67 O

ATOM 30820 N GLU B1141 22, .107 -32. .160 55. .353 1. .00 58. .35 N

ATOM 30822 CA GLU B1141 21. .300 -33. .375 55. .382 1. .00 56. .90 C

ATOM 30824 CB GLU B1141 21. .413 -34. .093 56. .739 1. .00 57. .44 C

ATOM 30827 CG GLU B1141 22. .740 -34. .841 56. .873 1. .00 59. .94 C

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VD t- vo in LO in in to LO m m LO in m in LO m LO in n m LO o o o o O o o o o o o o o o o O o o o o o O o o o o O o o o o o o o o o o o o o o o o O

H vi H rH H H vi rH vi H vi H rH vi - H v-i v- H i rH vi

H v-i ro H oo em CD o sμ CM co on rH vo v-i oo VD oo VD CM CM O o m oo cn 00 VD co sμ sμ 00 ro in on vo vo ro CM ro CM vi sμ tr- ro CM sμ o VD H I- CM o co ro on LO v-i IO sμ t- sμ 00 CM oo cm oo LO sμ LO sμ LO co I- cn CM CM rH o o o cm oo o cm in m in LO in LO in IO LO LO m LO LO in LO LO IO LO sμ sμ sμ sμ

LO CM vo VD sμ oo cn cm cm VO t- t- LO sμ CM CM ro VD m on in o en o t- LO o H CM in CM oo 10 sμ CD CM cn s ro vi o o to

H sμ CM o on ∞ sμ ro vo ro CM oo CM on lO LO m tr- o t- VD 00

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1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 r- sμ ro cn em I- cn VD I- tr- r- LO oo vo ro r- o vo CM o oo o vo vi on cm oo I- cn CM CM sμ t- s CM o O cn CM in sμ ro rH LO H cm t- ∞ vi sμ vi VO co CM m o O vi O CM v-i o sμ ro H oo ro CM oo cn D on oo t- t- oo 0 oo I- σn on o oo oo t- VD LO ro

CM CM CM vi vi rH H H H vi vi H rH H vi CM vi vi H vi H vi

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H vi vi vi vi vi v-i H vi v v-i H H vi i v-i vi rH H H v-i vi

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U U U U a U U U U U U U a U u U u a U U U o v-i CM ro sμ m I- on CM sμ oo CM cn sμ VD oo CM 00 sμ vo co vi cn ro 00 ΓO ro cn 00 en sμ sμ sμ m in LO LO LO vo VD vo VD vo r- co oo oo co oo oo 00 oo oo co oo oo oo oo oo 00 oo co oo oo co ∞

CD o o o o o o o o o o o o o o o o o o o o o cn ro oo ro on ro cn ro ro en ro 00 cn en 00 ro ro ro ro en cn ro

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 U U U U U U U U U U U U U U U U U U O O O O EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH ft fit fit fit fit fit fit ft fit ft fit fit ft ft fit fit fit fit ft fit fit fit u o o u u a u u u u u u u a u u u u u u a u

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LO LO vo in LO LO LO LO LO LO VD sμ sμ sμ LO sμ sμ sμ IO LO LO LO o o o o o o o o o o o o o o O o o σ o o o o o o o o o O o o o o o o o o O o o o o o o o

H rH rH vi vi H H v-i H v-i H H H H rH vi vi vi vi H i r-i

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H sμ o r- cm oo m CM cm o VD r- co sμ CM ro sμ VD vi CO ∞ 00 CO vo H i cm o tr- o VD in on oo o ∞ O oo CO CM 00 CO cn rH ro sμ sμ H σn CM H ro CM CM ΓM o i on cm r- r- σn rm on D H cn cn ro ro CM 00 en cn ro cn on ro cn CM CM CM CM CM CM CM cn ro

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

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H vi i H vi vi vi vi rH H v- i i H vi H H vi H H vi H

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P p p p p ffl ffl ffl ffl ft ffl P- ffl O O O O O O 0 fiϋ fiϋ P P P P CΛ W trΛ irΛ TO W ω ffl flf ffl ffl ffl ffl ffl P P

U ø ø o ø fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ ffl ffl ffl ft ffl f^"1, ffl fiϋ fiϋ v-i CM H CM u P PQ PQ f ffl CD P P fit ffl KD P ft

U U u O a U U U U U U O a U U U U U U a U sμ io VD I- oo on H ro VD I- oo on o i CM sμ iro ro sμ LO r- t- t- I- I- r- IT- oo oo oo ∞ oo oo em σn σn cm on o o o o o

∞ oo oo oo oo CO oo oo oo ∞ oo oo oo oo oo oo oo cm em em cm em o o o o o o o o o o o o o o o o o o o o o o en en ro en en on ro ro ro ro ro en en ro cn en ro ro ro cn cn ro

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 U U U O O O O O O O O O O O O O O O O O O O EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH ft ft ft ft ft fit ft fit fit fit fit fit fit fit ft fit ft fit fit ft ft fit u u u a u u u u u u u u u u u a u u u u a u

cm I- sμ o r- in vo IO sμ m CM co VD cm vo sμ CM LO o en t- vi

CM o oo oo r- t- LO sμ o o H CM VD vi on vi x-i I- sμ sμ vi vo sμ I- vo cm rH CM oo en m VD oo t- LO CM CM CM CM H CM H cn sμ

LO lO LO m VD vo vo t- L-^r I- I- t- t- VD vo vo IO ID VD VD VD vo o o o o O o o o o o o O o o O o O O o o o o o O o o O o o o o o o o o o o o O o o o o o vi H vi H rH vi vi H H H v-i i vi rH vi vi vi vi H i rH vi o CO en en cn vo vo r- tr- ∞ CM CM v-i VD o co CM cn ro cn oo VD

ID ro vo oo CD cn vi CM CM CM O VD sμ 00 tr- LO troo oo t~- I- sμ on vi CM oo CD vo t- ^■sμ sμ I- I- o o O CM tr- io oo H oo cm LO o en ro oo in LO IO VD t- 00 cn on oo VD D lO r- r- r- t- m LO sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ sμ o vi vo vi co vi H CM o CM ro oo LO ro on ro cm cn ro t- CM H cm t- CM I- co o sμ LO sμ en vi sμ sμ VD oo oo CM oo sμ ID oo ro o o o oo vo sμ ro VD in vi O oo on co on on o 00 LO ID o LO

CM CM CM CM ro sμ LO VD t- r- co LO sμ CM CM H vi cm O o o on ro ro ro ro ro cn ro ro ro en ro ro ro cn en ro cn CM ro ro en CM

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H on oo o o vi O o o D on o o em oo o on o oo I- oo vo v-i vi H vi H H vi vi vi vi vi vi

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H H

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CM H vi vi

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H vi vi vi H rH vi vi vi rH vi vi H vi H vi H H H H vi vi

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H H H H H H vi vi H H H vi H vi H vi H vi H H H H

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cm σn cm cm cm o o o o o o o O rH H H vi vi CM CM CM CM

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1 H H H H 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

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o o o o o o o o o o O o o o o o o o o o o o o o o o o o o o O o o o o o o o v-i H vi H v-i H vi H v-i H H H H H H H H H vi H vi H en H H H iron in I- ro cm cm sμ oo en on r- cm r- sμ on cm lO H oo H en sμ ro in I- ro o o oo oo on IO ro irsμ vi lO in cn ro ro ro ro cn ro sμ on sμ sμ t- irlO VD vo ID vo ID vo vo vo vo

CM o ΓM o ro ro CM vo oo cm o o o

1 H H 1 1 ro oo o sμ vo t- o VD m ro vo oo t- o ro oo 00 cn 00 H oo sμ cm CM I- o o in cm 00 o to cm oo oo vi o ro oo sμ t- ∞ ro H ro ro on m sμ 00 vi o cn ro

H vi H vi H H H H vi i i vi vi H

o o o CD CD i i vi vi H H vi vi H vi H CM CM CM CM CM CM cn cm rm cm on on CD CD on on em CD cm on on on cm cm cm on cm cm

H i H H H H H vi vi vi H H H H H H H H vi H H H vi vi H H i-i H H H H H H H vi H H H H H H H H vi

PQ ffl PQ ffl ffl PQ ffl ffl ffl ffl Q PQ ffl ffl ffl ffl ffl PQ Q ffl PQ ffl

0 0 0 0 0 PQ pq PQ PQ PQ pq pq pq pq pq ffl pq pq PQ pq pq PQ ffl ffl ffl ffl i Si S3 J SJ J SJ Si Si Si Si P P P P P P fiu fit fit fiϋ fit ffl ffl ffl ffl ffl ffl ffl ffl ft ft ft H H H H H H

H CM H H CM CM v-i vi CM

N i i fit ffl 0 o pq N pq P fiϋ ffl 0 Ω 0

0 a a U O a U U 0 U U U u U U u a U U u U U ro sμ I- o i CM sμ vo σn o CM sμ VD ∞ o H CM sμ VD oo H m

I- I- I- oo oo oo oo oo oo cm cm on on on o o o o o o H vi

LO LO m LO LO to LO to to m in m in in IO IO VD VD VO vo VD vo i H H H H H H r-i H H H vi H H H H H H H H H H en ro ro on ro on on 00 ro cn ro cn ro cn en on ro on on on 00 on

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2

U U U U U U U U U U U U U U U U u U U U u U

EH EH EH EH EH EH EH EH EH EH hi EH EH EH EH EH EH H EH EH EH EH ft fit fit fit fit fit fit fit ft fit fit fit ft f fit fit fit fiϋ fit ft fit ft

111 u u a u u u u u u a u u u u o o u u a u u u

tr- io sμ sμ H on 10 ro r- CM sμ cn vo sμ m t- vi m i oo o o

CM VD oo m to oo CM co lO CM o H irH cn LO O o sμ sμ oo cn

CM cn o o σn on i H o IO tr- tr- on in o oo r- vo troo oo on LO in LO LO sμ sμ m in LO LO LO LO IO VD I- vo LO m io in to LO o D O o o o o o o o o o o o o o O o o o o o o CD O o o o o o o o o O o o o o o o o o o o rH r-H H H H i H H H r-t H v-i i H vi H vi rH H H rH H o VD ID o σn t- on s-μ H cm t- CD σn cm vo oo rr- H CO 00 00 tr-

00 CM 00 co cm o CM LO 00 VD o VD on H vo oo ro VO CM in o on r- r- co o to cn oo t- vi o cm en CM in sμ LO LO iro on o CM

LO vo in VD LO sμ c- I- σn σn H CM en sμ oo σn on cn 00 on O ID vo D VD VD VD vo vo VD vo vo r- r- t- t- vo VD ID vo vo I-

H sμ sμ VD VD H vo 00 o in oo in m H t- o 00 VD o cm CM H o CM on σn in LO en ro -μ sμ m in ∞ r- on on CM CM t- vi in r- vo ro o oo on cn em r- oo IT- vo CM on sμ in H σn VD o μ cm σn cn O H H H D H CM H CM CM ro CM CM r-i ji sμ sμ VD ID to

1 1 H H rH i i H H i H H H -i i rH H rH rH H i

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 I I 1 1

CM CM sμ o o CM co sμ tr- CO cm I- r- H cm LO ro t- sμ t- τ-1 CM

H o sμ ro oo lO CM in i vo LO sμ on sμ cn H r- o sμ LO ro VO σn t- t- m H r- ir- ro CD on i t- t- o CM O o o on i^rCM VO o vi σn cn ∞ t- co o H o vi o H CM H 00 vi CM σn on oo I- vi v-i H H i vi vi H H H vi vi vi

CM CM oo on cn on ro ro on sμ sμ sμ sμ sμ sμ sμ sμ sμ LO to in LO on σn em σn cm cm en σn cm σn cn on cm en σn cm cn cm cm σn cm on

H H vi i H vi H H H H H H H H H H vi vi H H vi H

H H vi vi H vi H H H H H H H H H H vi H H H vi H ffl ffl PQ Pi ffl PQ ffl PQ PQ PQ ffl PQ PQ ffl ffl ffl PQ PQ PQ ffl Pi PQ

PQ PQ o O O o o o q P P P P P P P P P P P P P

P P ffl ffl ffl ffl ffl ffl ffl P P P P P P P P P fiϋ fiϋ fiϋ fi

H H ft ft ft ft ft ft ft ø 0 0 0 0 0 0 0 0 > > > >

CM fit PQ CO Ω fit ffl CO Q PQ PQ fit ffl CO u O a U U U U U a U U U U U U U O a U u U cm o H CM sμ tr- o ro sμ LO r- cm CM LO vo r- ∞ cm o CM sμ VD

H CM CM CM CM CM ro ro en 00 ro ro sμ sμ sμ sμ s-μ sμ IO in in LO vo 10 10 10 0 10 io vo VD VD VD vo VD VD vo VD o vo vo VD VD VD

H H H H H vi vi i H H H H H H H H vi H H H H vi on on 00 on on cn cn cn on cn on on ro ro ro ro cn on ro ro on en

2o o2 2o 2o 2o 2o 2o 2o o2 2o 2o 2o 2o 2o 2o o2 2o o2 o2 2o o2 2o

EH EH H EH EH EH EH EH EH EH B EH EH EH EH EH EH EH EH EH EH EH ft ft ft ft ft fit fit ft ft ft fit fit fit ft ft fit ft fit fit fit ft ft ø ø o a ø u u u a u u u u u u u u a u u u a

m ro o sμ en o vo sμ ∞ vi 10 o co 00 o sμ I- o oo VD on sμ oo o I- o ro co en sμ I- sμ H oo cm vo in cn t> on c- c- oo vi oo vo o o m m to sμ sμ sμ ro ro ro s en en in vo vo o o o o o o o o o o o o o o o O o o O o o o o o o o o o o o o o o o o o o o O o o

H rH rH rH r-i vi vi ^•H H H H r-i rH ^•H rH rH v v- vi vi rH H o t- co co ^■sμ vi t- VD r- on sμ O LO sμ oo oo ro sji o sμ t- vo v-i ω o ro VD tr- co o tro r- I— ro vo o ro LO ∞ o vo I- t- cn io trlO sμ sμ sμ sμ ro 00 LO sμ vo oo cm vo io ID VD vo vo vo vo VD vo vo vo VD vo vo vo VD oo o t- vo sμ LO ' r- LO H cm sμ LO H to on ro oo oo 00 co ro sμ o 00 00 LO t- o i^rsμ vi on

∞ H H IO I- CO H 00 t- n vo on in in ID in in VD in sμ ro on sμ LO LO sμ ro vi i H H -i H H H i v- H H H H vi vi 1 1 1 1 1 1 1 1 1 1 1 1 ro ro sμ cn sμ H r- σn L— vo vo sμ O cm oo VD vi oo em ro sμ ro H sμ H oo t- o vo on oo o I- o 00 ro cn ∞ o VD o t- r- t- sμ H oo O H I- on t- o on ri on on t- I- oo ∞ C- VD vo on cn cn in LO sμ sμ

H H vi vi i H H H vi

in LO LO vo vo VD vo VD vo VD VD vo vo vo VD VD VD I- irtr- irI- cm cm cm σn cm on cn on cm on on on on cn cn on σn cm on on on cn vi vi H H H vi H vi vi H H vi H H H vi H H vi vi v H vi vi H H H H H vi H H H vi H H H H H H vi vi vi H ffl ffl ffl ffl PQ PQ ffl ffl ffl ffl ffl ffl ffl ffl m PQ ffl PQ ffl ffl ffl PQ p p p ft ft ft ft ft ft ft ft ft ft ft ft ft ft co ϋ- i- co co fiϋ fiϋ fiϋ ffl ffl ffl ffl ffl ffl PH Pπ ffl ffl ffl ffl ffl ffl H H H H H > EH EH EH EH EH EH EH EH EH EH EH EH EH EH SI SJ S; SJ :

CM H H CM CM cn on CM CM H O fiϋ ffl CO P pq pq o PQ N SJ IS] fit ffl CO P u U u a U U U U a U u U U U u u u a U U U a o sμ LO VD co o 00 sμ ID CO cm o CM sμ VD oo on o CM sμ irco

ID vo VD VD vo t- trtrI- I- t- co oo oo oo co co cm cm on on on

10 VD VD VD vo VD io io ID D ID vo vo vo vo vo VD vo vo vo VD VD

H H H H H H vi vi vi H H H H H H H H vi H H H H ro ro ro on on ro cn cn en ro en on cn ro on 00 on cn ro ro ro ro

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 U U U U U U U U U O O O O O O O O O O O O O EH EH EH EH H EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH ft fit fit fit fit fit fit fit fit fit fit fit fit fit fit ft fit fit fit fit fit fit u a u u u u a u a a a u a a cf- u u u o co o o

CM LO sμ CO LO oo o oo sμ sμ CM H vo r- on VD oo ro in ro ID cm H o o σn ro on vo VD o on oo ro oo r- H o on H c- CM on ro en LO vi o CM H CM on ro on H sμ LO σn CO r- CM co LO to in

VD vo D ID vo vo ro ro CM ro CM ro cn CM co ∞ co 00 co o o o

H H vi o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o

H vi H H vi rH H r-i rH rH i rH rH rH H vi vi H vi rH H vi

VD on CM r- co o ro cn LO em LO on ro r- r- H VD r- VD o CM LO sμ cn cm ro ro m I- o o sμ sμ in vi H o sμ on o t- H LO vo cn vi ID H o H LO CM cn r- ro CM vo vo in H σn in m sμ 00 LO

CO I- t- vo oo CM CM CM H CM vi o o CM v- CM CM m tr- oo VD vo ID VD vo VD CM CM CM CM 1 1 1 1 CM CM CM CM CM CM C CM trH H LO ro v-i VD cn vo o ∞ vo o ro vo on oo ir- o oo O CM io sμ H CM VD sμ O o VD cm LO r- in on CM on sμ co H H LO cn

H CM LO r- o sμ oo o o H o ro LO VD co t- sμ LO sμ cn cm sμ vi H CM vo t- t- cn H CM vi LO in vo sμ o σ on H H vi o H vi H H H H H ro sμ sμ sμ v-i H H vi cn en CM ro cn vi H H

1 1 1 I 1 1 1 1 1 1 1 1 1 1 1 1 1 1 sμ cm sμ CM H VD cm cm I- in co o I- on s ID co s-μ vi H v- r- in cm m m σn LO CM VD sμ oo co on lO oo t- H em o vo en oo vo sμ iron IT- H on vo oo o o oo in ro LO LO vi CM ro VD LO co vo sμ on ro LO vo LO sμ sμ sμ vo troo oo on CM cn CM H cn sμ m ro vi vi vi v-i H v-i sμ sμ sμ sμ io LO LO in IO m m in m ro ro ro

I- t- irr- t- t- on en on cm o o O o H vi H H H CM CM CM σn σn on σn σn cn oo 00 oo ∞ cm on on cm em en em cm on cn cm cm

H H H H H H H H vi H H H H H H vi vi H H H vi H

H H H H H H ffl Q ffl ffl ffl PQ rs 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2

CO O) co CO CO CO H H H H H H H H sμ sμ sμ sμ sμ sμ sμ sμ

H H H H H H fiϋ fit fiϋ fiϋ fiϋ fit, fiϋ fiϋ O o O O O O O O

SJ SJ Si i SJ Si 0 0 0 0 0 KD 0 0 co co CO co CO O CO CΛ

H

a u ro CM H ro CM H CM 00 sμ v CM a a a U a a CO o o U O C/3 O O o CM sμ vo I- co CD H CM LO oo o H sμ r- oo cm o H CM cn sμ o o o o o o CD H H H vi CM CM CM CM CM CM ro ro ro cn ro

I- I- t- t- I- t- I- r- I- I- I- I- I- r- r- I- r- r- t- t- t- t-

H H H H H H H H H H H H H H vi H H H H H H H ro ro ro en on ro 00 ro on ro ro ro ro 00 en ro ro ro on ro ro ro

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 O O O O O O O O O O O O O O O O O O O O O O EH EH EH EH EH EH EH EH EH EH EH EH EH EH EH E^H E^H E^H E^H E^H EH EH fi fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ ι fiϋ fiϋ fi fi fiϋ fiϋ fiϋ fi fiϋ fi fi fiϋ fiϋ o o o o o o o o o o o o o o o o o o o o o o

o s cn VD vi o ro t- vo CM I- LO VD CM oo H trVD ID LO CO CM to VD CD in CD in vi CM cm CM oo 00 o on oo O io H 00 00 r-t ri sμ tr- CD VD VD i^rVD o LO on sμ CO VD σn to H VD LO on on CM L0 o o cn sμ ro on sμ m en sμ sμ in in sμ in LO ro LO IO sμ vi H CD O CD o o o o o o o o O o o o o o o o o O o o O CD o o o o o o o o O o o o o o o o o O o vi H H H H H vi H vi vi H v-i vi i H H vi v-i H v-i H - ir- CD o ro r- CM cn o i CM H CM v-i CD en ro cn I- H CD O v-i ro cn LO LO I- en CD VD cm r- sμ O en σn cn sμ o tr- VD cn VD I-

VD VD cm o CM vi r- CM sμ 10 CM VO vo oo LO H o co CM LO on H vo VD cm cn en vo sμ VD co I- O VD en LO oo O sμ H LO en CM cm ΓM CM CM 1 H H CM vi CM ro vi 1 1 vi 1 LO sμ i cm ιn oo em r- vo LO LO -i 00 CM ∞ vo LO VD sμ sμ en CM ro vo VD H H sμ sμ CM r- t- vi vi ID oo oo CM vi ro r- oo LO o 00 in in CM ro t- LO on H r- H VD ro sμ o LO vi H vi o CM in I- vi

CM o CM CM o sμ oo H CM r- sμ H cm CM t- t— o cm o r- i vi H ro VD t- CM ro CM CM VO 10 VD s-μ VD 1 ro in sμ CM I- vo cn 1 1 1 1 1 sμ o cm CM oo o H cn sμ LO cn CM ro VD CM on in CM oo sμ r- cn cn sμ o en σn VD ro sμ LO LO cn o on i CM cn ID oo ID m o m cm r- CD σn H sμ sμ t- CM m VD ro co LO vi sμ on CM H en 00 sμ en ID cn CM CM t- vo o vi CM sμ ro H o VD cn vi CM VO cm ro cn LO LO m sμ sμ ro r- sμ 1 tr- vo I- sμ in CM LO VD ro

CM CM cn sμ m VoO tt-- oooo aDn o CD H H C CMM rcno sμ LO vo t- oo en o vi CM em on cm cm cm ccmm σσnn oonn c CmD o CD oo o O o CD o o o o o o H vi H vi H vi H vi v vii H v- vii v Hi C CMM C CMM C CMM C CMM CM CM CM CM CM CM CM CM CM

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2

ro sμ

O U u o o o o o o o o o o o o o o o o o o o

LO vo t- O cn VD on CM LO oo vi s-μ t- o on vo on ΓM m 00 vi ro cn an sμ s-μ sμ sμ LO LO to vo vo ID I- c- r- t- oo co oo on cm rr- r- r- t- r- tr- t- t- t- t- rr- rr- tr- I- r- t- r- t- rr- r- r- t- vi H H H vi H H vi H H H H i H H H H vi r-i H H H cn on en cn en ro ro en ro ro cn 00 cn 00 ro 00 ro cn 00 on ro on

2o 2o 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 o o o o 2 o o O o o o o U u U U U u u O

H EH EoH o H^" H H H H H H H H H H H H H H H H H H f fit fit fit f fiϋ fiϋ fiϋ fiϋ fit fiϋ fiϋ fiϋ fit fit fiϋ fiϋ O O O O O O O O O O O O O O O O O O O O O O

CM ro H ir00 to ro sμ on ro ro oo I- vo o sμ LO sμ VD cm vo oo sμ t- t- on ro 00 in r- vo o t- sμ ro o o sμ CM sμ I- H VD CM r- sμ CM to I- VD o H in vo r- m VD LO ro t- sμ CM o vo LO VO

LO LO in LO LO LO VD sμ vo sμ sμ in in LO vo lO LO vo VD LO vo VD o o o o O o o o o o o o o o o o o o o o o O o o o o O o o o o o o o o o o o o o o o o O

H H vi H H H H H H H H vi i v-i H H vi v-i H v-i vi H vo o LO o H on o oo CM sμ VO CM en CD CM sμ LO cn on t- H 00

CM VD ro H H o on vi σn cn sμ LO 00 LO IO in 00 m r- m H CM σn LO LO CM μ ro t- sμ ro sμ sμ VD sμ sμ 00 10 c- H CM vo O sμ vo H CM cm sμ sμ o sμ H sμ in ΓM vi vo H CM lO co o VD on i^rro H i i CM LO H CM in sμ sμ s vi r-i 1 v-i in H CM VD vo 1 1

LO 00 oo 00 H sμ o cm oo H ro on oo LO H CM oo vi oo ro co on oo VO cm ro sμ cm irt- VD CM ro vi sμ sμ sμ v-i sμ en CM ro vo I-

I- t- o μ sμ t- on vo o I- ro CD en CM co v-i oo CD VD o 00 sμ sμ IΩ vi ID o CM vo CM oo o in cn CM H I- H o LO m cm H VD vo sμ sμ sμ LO VD t- 1 t- sμ ro CM I- vi 1 vi CM

1 1 1 1 1 r- CM co o ro H 10 on CM oo o cn H oo ro m CM CM v-i ro co sμ

CM en o LO r-i ro in VD I- tr- cn sμ vo sμ vo LO CM o CM o vo in m t- o o ID to oo o sμ co H L-^r co vo CM o r-i VD on I- H t- on sμ oo I- vi VD cn H on o cm oo I- rr- VD o ID CM o r- sμ ∞ sμ ro CM m LO sμ sμ CM VD sμ LO CM m CM H CM CM

cn sμ LO VD I- co cm O vi CM ro sμ LO vo t- oo cm o -i CM ro sμ

H vi H H H H H CM CM CM CM CM CM CM CM CM CM ro en ro ro ro

CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM CM

2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2

o o o o o o o o o o o o o o o o o o o o o o r- o en vo on CM LO co H sμ t- O cn VD cm CM LO oo vi sμ t- o cn o o o o vi vi vi CM CM CM en en cn ro sμ sμ sμ in m LO vo t- oo co co co co co co oo co ∞ oo oo co ∞ oo co co oo oo CO oo vi H i H r-H vi H H vi vi H v H H vi vi vi H H r-t H H en ro cn ro ro en cn ro cn cn 00 cn ro ro cn cn cn ro on cn on ro

2o 2o 2o 2o 2o 2 2 o2 2 2 2 2 2 H oEH oEH EoH oEH o 2 2 2 EH oEH oEH o 2 EH o 2 2 2 2 2

EH EH EH EH EH oEH oEH E EH oEH oEH oEH oEH oEH f^ f^ ft ft fit fit fit ^fit fit fit f fit fi fit fit ^ft fit fit fit ft fit ft O O O O O O O O O

H ro cn oo H cn H ro o sμ LO ro LO oo CM oo on CM

o o o o O o o o o o o o o O o o o o

H H H H H H H H H

00 LO VD on on o s H t- sμ ID vi o ro VD H IrCM ro 00 sμ t- r- I- VD an cm

ro ID cn CM VD O O LO LO vi CM H LO LO cm O H LO cn sμ on VD CM r-i O sμ 00 sμ O I- sμ CM H VD CM oo

H ro H r- I- I- H CM

1 1 1 sμ o oo O ID σn o sμ CM

I- o I- VD ΓM ro VD CM o CM H σn LO I- ro sμ vo CM

to VD I- CO cn o H CM ro ro ro ro oo cn sμ s sμ sμ

CM CM CM CM CM CM CM CM CM

2 2 2 2 2 2 2 2 2

o o o o o o o o o

fiϋ fi fi fiϋ fiϋ fiϋ fiϋ fiϋ fiϋ

Claims

What is claimed is:

1. A crystal of Herpes simplex virus 1 DNA polymerase having orthorhombic space group symmetry ¥2{λ{2_\ and having a unit cell defined by the dimensions a, b, c, , β, and γ, wherein a is about 80 A to about 130 A, b is about 100 A to about 150 A, c is about 200 A to about 250 A, and α=β=γ=90°.

2. A method for crystallizing a Herpes simplex virus 1 DNA polymerase molecule or molecular complex comprising: providing a solution comprising purified Herpes simplex virus 1 DNA polymerase and guanidine-HCl; and inducing Herpes simplex virus 1 DNA to crystallize from the solution.

3. A Herpes simplex virus 1 DNA polymerase having an amino acid sequence comprising SEQ IDNO: 1.

4. A Heipes simplex virus 1 DNA polymerase having an amino acid sequence consisting of SEQ IDNO:l.

5. A method of preparing a Herpes simplex virus 1 DNA polymerase comprising digesting a fusion protein constructed from a maltose binding protein and full length Herpes simplex virus 1 DNA polymerase.

6. A method of identifying the ability for a potential modifier to bind to Herpes simplex virus 1 DNA polymerase, the method comprising: exposing a crystal of Heφes simplex virus 1 DNA polymerase to one or more samples comprising a potential modifier of Herpes simplex virus 1 DNA polymerase; and determining whether a potential modifier-Heφes simplex virus 1 DNA polymerase molecular complex is formed.

7. A method of acquiring structural information for designing potential modifiers for forming molecular complexes with Heφes simplex virus 1 DNA polymerase, the method comprising: exposing a crystal of Heφes simplex virus 1 DNA polymerase to a library of potential modifiers having diverse shapes; and determining whether a potential modifier-Heφes simplex virus 1 DNA polymerase molecular complex is formed.

8. A molecule or molecular complex comprising at least a portion of a Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active site, wherein the active site comprises the amino acids listed in Table 2 and the active site is defined by a set of points having a root mean square deviation of less than about 3 A from points representing the backbone atoms of said amino acids as represented by the stracture coordinates listed in Table 1.

9. A method for obtaining structural information about a molecule or a molecular complex of unknown structure comprising: crystallizing the molecule or molecular complex; generating an x-ray diffraction pattern from the crystallized molecule or molecular complex; and applying to the x-ray diffraction pattern at least a portion of the structure coordinates as set forth in Table 1 for Heφes simplex virus 1 DNA polymerase to generate a three-dimensional electron density map of at least a portion of the molecule or molecular complex whose structure is unknown.

10. A method for homology modeling a Heφes simplex virus 1 DNA polymerase homolog comprising: aligning the amino acid sequence of a Heφes simplex virus 1 DNA polymerase homolog with an amino acid sequence of Heφes simplex virus 1 DNA polymerase and incoφorating the sequence of the Heφes simplex virus 1 DNA polymerase homolog into a model of Heφes simplex virus 1 DNA polymerase formed from structure coordinates as set forth in Table 1 for Heφes simplex virus 1 DNA polymerase to yield a preliminary model of the Heφes simplex virus 1 DNA polymerase homolog; subjecting the preliminary model to energy minimization to yield an energy minimized model; and remodeling regions of the energy minimized model where stereochemistry restraints are violated to yield a final model of the Heφes simplex virus 1 DNA polymerase homolog.

11. A computer-assisted method for identifying a potential modifier of Heφes simplex virus 1 DNA polymerase activity comprising: supplying a computer modeling application with a set of structure coordinates of a molecule or molecular complex comprising at least a portion of a Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active site, wherein the active site comprises the amino acids listed in Table 2 and is defined by a set of points having a root mean square deviation of less than about 3 A from points representing the backbone atoms of said amino acids as represented by structure coordinates listed in Table 1 ; supplying the computer modeling application with a set of structure coordinates of a chemical entity; and determining whether the chemical entity is expected to bind to or interfere with the molecule or molecular complex, wherein binding to or interfering with the molecule or molecular complex is indicative of potential modification of Heφes simplex virus 1 DNA polymerase activity.

12. A computer-assisted method for designing a potential modifier of Heφes simplex virus 1 DNA polymerase activity comprising: supplying a computer modeling application with a set of stracture coordinates of a molecule or molecular complex comprising at least a portion of a Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active site, wherein the active site comprises the amino acids listed in Table 2 and is defined by a set of points having a root mean square deviation of less than about 3 A from points representing the backbone atoms of said amino acids as represented by structure coordinates listed in Table 1; supplying the computer modeling application with a set of structure coordinates for a chemical entity; evaluating the potential binding interactions between the chemical entity and active site of the molecule or molecular complex; structurally modifying the chemical entity to yield a set of structure coordinates for a modified chemical entity; and determining whether the modified chemical entity is expected to bind to or interfere with the molecule or molecular complex, wherein binding to or interfering with the molecule or molecular complex is indicative of potential modification of Heφes simplex virus 1 DNA polymerase activity.

13. A computer- assisted method for designing a potential modifier of Heφes simplex virus 1 DNA polymerase activity de novo comprising: supplying a computer modeling application with a set of structure coordinates of a molecule or molecular complex comprising at least a portion of a Heφes simplex virus 1 DNA polymerase or Heφes simplex virus 1 DNA polymerase-like active site, wherein the active site comprises the amino acids listed in Table 2 and is defined by a set of points having a root mean square deviation of less than about 3 A from points representing the backbone atoms of said amino acids as represented by structure coordinates listed in Table 1 ; forming a chemical entity represented by set of structure coordinates; and determining whether the chemical entity is expected to bind to or interfere with the molecule or molecular complex, wherein binding to or interfering with the molecule or molecular complex is indicative of potential modification of Heφes simplex virus 1 DNA polymerase activity.

14. A method of using the crystal of claim 1 in a drag screening assay comprising: selecting a potential modifier by performing rational drag design with a three-dimensional structure determined for the crystal, wherein selecting is performed in conjunction with computer modeling; contacting the potential modifier with an active site of Heφes simplex viras 1 DNA polymerase; and detecting an affinity for binding of the potential modifier with the active site, wherein a potential drag is selected on the basis of its having greater affinity for the active site of Heφes simplex viras 1 DNA polymerase than that of a known modifier for the active site of Heφes simplex viras 1 DNA polymerase.

15. A method of using the crystal of claim 1 in a drag screening assay comprising: selecting a potential modifier by performing rational drag design with a three-dimensional stracture determined for the crystal, wherein selecting is performed in conjunction with computer modeling; adding the potential modifier to a binding assay; and detecting a measure of binding, wherein a potential modifier that binds is selected as a potential drug.