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Clamp loader ATPases and the evolution of DNA replication machinery
Clamp loaders are pentameric ATPases of the AAA+ family that operate to ensure processive DNA replication. They do so by loading onto DNA the ring-shaped sliding clamps that tether the polymerase to the DNA. Structural and biochemical analysis of clamp loaders has shown how, despite differences in composition across different branches of life, all clamp loaders undergo the same concerted conformational transformations, which generate a binding surface for the open clamp and an internal spiral chamber into which the DNA at the replication fork can slide, triggering ATP hydrolysis, release of the clamp loader, and closure of the clamp round the DNA. We review here the current understanding of the clamp loader mechanism and discuss the implications of the differences between clamp loaders from the different branches of life.
High-speed replication of chromosomal DNA requires the DNA polymerase to be attached to a sliding clamp (known as proliferating cell nuclear antigen, or PCNA, in eukaryotes) that prevents the polymerase from falling off DNA
Because sliding clamps encircle DNA but do not interact tightly with it, they can slide along the double helix by diffusion
The increase in the processivity and speed of DNA synthesis when DNA polymerases are engaged to sliding clamps is very considerable. For example, in the absence of the clamp, the polymerase subunit of the bacterial replicase synthesizes DNA at a rate of about 10 base pairs per second
The enhancement in speed and processivity that the clamp confers on the polymerase would not be possible without the clamp loader, the less glamorous but much more hardworking handmaiden of the sliding clamp, which diligently loads the clamps onto primed DNA throughout the process of DNA replication. Together, the clamp and the clamp loader lie at the heart of the replisome - the DNA replication machinery, which, with the polymerases (leading and lagging strand), includes the helicases that unwind the double-stranded DNA ahead of the polymerase at the replication fork, the primase that synthesizes the RNA primer required for the initiation of DNA synthesis, and the single-strand DNA-binding proteins that prevent the DNA from re-annealing in the wake of the helicases (Figure
Figure 1
Architecture of the bacterial replication fork
Architecture of the bacterial replication fork. The helicase is a homohexamer that encircles the lagging strand and binds directly to the primase synthesizing the primer RNA. The clamp loader acts to hold the replisome together by binding directly to the helicase as well as three polymerase subunits for simultaneous synthesis of the leading and lagging strands. The leading strand polymerase synthesizes DNA continuously, while the other two polymerases presumably cycle on and off the lagging strand, which is coated in single-strand binding protein (SSB). The polymerase subunits are attached to circular clamps that encircle duplex DNA for enhanced processivity and speed.
Despite the uncertainty in the precise architectural role of the clamp loader in the eukaryotic replisome, it is clear that sliding clamps are centrally important. The sliding clamp recruits the polymerase as well as other factors to the replication fork, including the chromatin-modifying proteins required to reassemble chromatin on the newly synthesized DNA
The clamp loader is a molecular switch operated by binding and hydrolysis of ATP
Clamp loaders are members of the AAA+ (ATPases associated with various cellular activities) family of ATPases
Initially thought to be a motor
Figure 2
Clamp loaders place sliding clamps at primer-template junctions for processive DNA replication
Clamp loaders place sliding clamps at primer-template junctions for processive DNA replication. When bound to ATP, clamp loaders are competent to bind and open the sliding clamp protein. This ternary complex can now bind to a primer-template junction, which activates the ATPase activity of the clamp loader. ATP hydrolysis causes the clamp loader to dissociate from the clamp and DNA, resulting in a loaded clamp that is competent for acting as a processivity factor for DNA polymerase. Figure adapted from
This complex but fundamental mechanism is embodied in an assembly of surprisingly diverse composition from bacteria to eukaryotes.
The structure of the clamp loader is more conserved than its composition
Unlike other AAA+ ATPases, which are typically hexameric, all clamp loaders are composed of five subunits in a circular arrangement, with a gap between the first and the fifth subunit, at the position of the missing sixth subunit. The individual subunits are designated A through E, starting with the subunit at the open interface and proceeding counter-clockwise around the clamp loader in the standard view (Figure
Figure 3
Composition of clamp loaders from the different branches of life
Composition of clamp loaders from the different branches of life. (a) Bacterial clamp loaders consist of three different proteins: δ, δ', and the τ or γ protein (γ, a truncation of the τ protein, is shown here). The δ protein is at the A position, with three copies of the ATPase subunits τ or γ at the B, C and D positions. The δ' protein sits at the E position. (b) Eukaryotic clamp loaders consist of five distinct proteins, RFC1 through RFC5. RFC1 lies at the A position, RFC4 at B, RFC3 at C, RFC2 at D and RFC5 at the E position. Eukaryotic A subunits contain an additional domain that bridges the gap between the A and E positions. (c) Bacteriophage clamp loaders consist of two distinct proteins. The gp62 protein, which lacks a AAA+ module but has an A' domain similar to that of eukaryotic clamp loaders, lies at the A position. The ATPase gp44 protein lies at the B, C, D, and E positions. Archaeal clamp loaders have a similar composition (see text for details). Figure adapted from
The clamp loader shares an essential aspect of its mechanism with other AAA+ complexes and other oligomeric ATPases, such as the F1-ATPase
Because clamp loaders are so fundamental to the replication process, it is no surprise that their structure and mechanism turn out to be highly conserved in all branches of life. This conservation has been somewhat difficult to appreciate, because the extensive biochemical analyses of bacterial, eukaryotic and bacteriophage clamps and clamp loaders have to a great extent proceeded independently in the past, and scientists working on these systems have used different conventions to identify the subunits. Adding to the confusion is the fact that, even though all clamp loaders have five essential subunits, the protein stoichiometry is different in clamp loaders from different branches of life. This confusion can be alleviated by using the simple A through E scheme for identifying the clamp loader subunits. Figure
The bacterial clamp loader is formed from three essential subunits: δ, δ', and the τ or γ protein (Figures
Figure 4
Structures of clamp loaders from different branches of life and in different bound states
Structures of clamp loaders from different branches of life and in different bound states. The five subunits of the clamp loader - A, B, C, D and E - are shown in different colors. (a) The structure of the clamp loader of
The eukaryotic clamp loader, Replication Factor C or RFC
The bacteriophage and archaeal clamp loaders are both composed of two proteins each with one unique protein occupying the A position (gp62 in T4 bacteriophage and RFC-l in archaea) and identical ATPase subunits at the B, C, D and E positions (the gp44 protein in T4 bacteriophage and RFC-s in archaea)
The bacteriophage clamp loader, whose structure seems to reflect a rather curious evolutionary history (we return to this briefly later), has played a particularly important part in the elucidation of the clamp loader mechanism. A recently determined structure of the T4 bacteriophage clamp loader bound to an ATP analog and in complex with the sliding clamp and primer template DNA
Recognition of the clamp during loading onto DNA
In three crystal structures of ATP-bound clamp loaders, the AAA+ modules can be seen to form a right-handed spiral
The right-handed spiral of the open clamp roughly matches the helical symmetry of DNA, with the clamp tracking along the minor groove of the DNA duplex. The right-handed spiral of the clamp can be described as a series of rigid-body rotations of the six domains present in the clamp (Figure
Figure 5
Clamp loader interactions to open the sliding clamp
Clamp loader interactions to open the sliding clamp. (a) The clamp is held in a distorted conformation. In the top panel, displacement vectors compared from the closed planar structure (PDB code 1CZD)
The recent structures of the T4 clamp loader bound to an open clamp and primer template DNA indicate how loaders open a clamp
The B, C, D and E subunits of the T4 clamp loader are identical (see above), yet they can occupy two very different binding surfaces on the clamp: the canonical binding cleft or the inter-subunit crevice (Figure
In both the T4 and the
The clamp loader, as we have seen, interacts with the clamp through the cooperative action of multiple interactions that appear to be weak. Most other clamp-binding proteins rely primarily on one tight interaction with the clamp. For example, most DNA polymerases
While the recently determined structures show how the clamp is held in the open state by the clamp loader, they do not address whether the clamp loader actively opens the clamp or simply traps a transiently open state. The T4 clamp appears to be dynamic and can open and close even in the absence of the loader
The clamp loader and the clamp collaborate in specific binding of the clamp loader to primer-template junctions
The clamp loader must position the sliding clamp on the DNA specifically at the primer-template junction where the polymerase is to be recruited. How the clamp loader recognizes the primer-template junction was first suggested by the structure of yeast clamp loader RFC bound to PCNA in the absence of DNA
Figure 6
A detailed model for clamp loading mechanism
A detailed model for clamp loading mechanism. (1) Prior to ATP binding, the clamp loader AAA+ modules are not organized in a manner competent to bind the clamp. (2) Upon fully binding ATP, the clamp loader AAA+ modules assume a spiral shape that can hold the clamp in an open lockwasher conformation. (3) This binary complex is competent to bind to primer template DNA. We propose that the dimensions of the open clamp restrict DNA access to the central chamber such that only single-stranded regions or a major groove at a single-stranded/double-stranded junction can slip through the crack in the clamp. (4) The duplex region of primer-template DNA then slides up into the clamp loader central chamber, which activates the ATPase. (5) ATP hydrolysis or Pi release initiates at the end of the AAA+ spiral, which allows the clamp to close. Further hydrolysis events cause release of the clamp loader from the clamp and DNA, resulting in a loaded clamp. Figure adapted from
The clamp loader AAA+ spiral does not conform to B-form DNA, but matches instead the helical symmetry of duplex A-form DNA
For DNA to bind in the central chamber of the clamp loader, it must enter through the clamp opening as well as the gap in the clamp loader spiral, which in the T4 clamp loader is flanked by the A' and A domains of the A subunit. The structures suggest, however, that both gaps are too narrow for either duplex DNA or a DNA-RNA hybrid to pass through. The clamp is only open by approximately 9 Å, and the gap between the A and A' domains of the clamp loader is as small as 14 Å
To explain how the DNA-RNA hybrid at the primer-template junction gains access to the chamber of the clamp loader, we propose that single-stranded DNA, or possibly the major groove of duplex DNA directly adjacent to a primer-template junction, initially enters through the opening in the clamp (Figure
DNA-triggered ATP hydrolysis drives ejection of the clamp loader
In all clamp loaders tested thus far, ATPase activity is greatly enhanced by the binding of primer-template DNA
The structural data suggest how DNA binding could play a role in switching on the ATPase activity of the clamp loader subunits. Earlier structural analyses of the bacterial and eukaryotic clamp loaders showed how the formation of tight intersubunit interfaces on ATP binding organizes the arginine finger at the catalytic center
In addition to the state fully bound by ATP, a ternary complex of T4 clamp loader, DNA, and clamp has been crystallized in a state in which the clamp loader B subunit is bound to ADP while the other active sites are bound to an ATP analog
The AAA+ spiral probably comes apart in a cooperative fashion, starting from one end of the spiral (Figure
One apparently consistent feature of clamp loaders from different domains of life is that only three of the ATPase subunits are catalytically active in clamp loading. (Mutated clamp loader complexes with fewer than three active ATPase sites can accomplish clamp loading
The T4 bacteriophage replication system seems to be a chimera, with functional modules borrowed from bacteria and eukaryotes
The genome of T4 bacteriophage is a curiosity, in that it is thought to be composed of genes derived from both eukaryotic and bacterial sources
Figure 7
Sequence and structural relationships between the T4 clamp loader and those from other branches of life
Sequence and structural relationships between the T4 clamp loader and those from other branches of life. (a) A sequence similarity dendrogram for the structurally characterized clamp loader subunits. The T4 clamp loader is more similar to yeast clamp loader than to the bacterial clamp loader subunits. Tree calculated using PHYLIP
One of the unique features of both the eukaryotic and T4 clamp loaders is the A subunit. The T4 A subunit forms an inverted U shape and reaches across the clamp interface
These findings may bear on the question of how the oligomeric character of the clamp influences the structure and mechanism of its loader. In particular, the trimeric clamps of T4 and eukaryotes are less stable
The similarity between the eukaryotic and T4 bacteriophage clamp loaders may reflect a deeper shared evolutionary history. The T4 DNA polymerase belongs to the B family, as do the eukaryotic replicases
These results imply that replisomes can be pieced together from different modules. The helicase, primase and single-stranded binding protein of T4 are most closely related to those of bacteria, while the clamp, clamp loader and polymerase are similar to those of eukaryotes. These two groups of proteins represent different functional modules of the replisome: the helicase and primase travel on the lagging strand as one unit, while the polymerase and clamp are tightly associated. These observations suggest that entire modules may be evolutionarily exchanged more readily than individual proteins.
Open questions
We have now reached a satisfactory state of understanding regarding the structural basis for clamp loader action, at least at the level of the general mechanism, understood in broad strokes (Figure
Although the inferences drawn from these structures are compelling, it is important to recognize that we only have one structure for some of the key steps in the clamp loading cycle, and that these structures are for clamp loaders that are very divergent in sequence. The structure of the ATP-free clamp loader is for the
Although we have emphasized the clamp loading aspect of the clamp loader machine, clamp loaders are critical for the proper coordination of leading and lagging strand synthesis. This is most clearly evident in bacteria, where the clamp loaders are physically attached to the polymerase subunits that replicate the leading strand continuously and those that cycle between Okazaki fragments on the lagging strand. The greater challenge ahead is to understand how clamp loaders serve to orchestrate rapid replication of chromosomal DNA, and the bacterial replisome is likely to provide the route to clearer structural understanding, because of the depth of biochemical information available, as well as the ability to purify intact complexes in reasonable amounts. We look forward to future advances in the structural analysis of intact replisome assemblies.
Additional file 1
The structure of the clamp loader of
Click here for file
Additional file 2
The structure of the budding yeast clamp loader, replication factor-C (RFC), bound to the sliding clamp, PCNA, and an ATP analog (PDB code 1SXJ).
Click here for file
Additional file 3
Movie of the T4 bacteriophage clamp loader bound to primer-template DNA, an ATP analog, and the sliding clamp during clamp closure (PDB code 3U60).
Click here for file
Insightful comments and guidance provided by Miranda Robertson and Stephen Harrison are warmly appreciated. We thank Kate Engel and Dr Tiago Barros for help with figure preparation. This work was supported by a Ruth L Kirschstein National Research Service Award through the National Institute of General Medical Sciences to BAK and by grants from the NIH to JK (GM45547) and MOD (GM38839).
Replisome-mediated DNA replication
Replisome mechanics: insights into a twin DNA polymerase machine
The diverse spectrum of sliding clamp interacting proteins
Loading clamps for DNA replication and repair
Motion of a DNA sliding clamp observed by single molecule fluorescence spectroscopy
Mechanism of the sliding beta-clamp of DNA polymerase III holoenzyme
Structure of a sliding clamp on DNA
Analysis of the role of PCNA-DNA contacts during clamp loading
Three-dimensional structure of the beta subunit of E. coli DNA polymerase III holoenzyme: a sliding DNA clamp
Structural and enzymatic studies of the T4 DNA replication system. I. Physical characterization of the polymerase accessory protein complex
Building a replisome from interacting pieces: sliding clamp complexed to a peptide from DNA polymerase and a polymerase editing complex
Crystal structure of the DNA polymerase processivity factor of T4 bacteriophage
Crystal structure of the eukaryotic DNA polymerase processivity factor PCNA
Crystal structure of an archaeal DNA sliding clamp: proliferating cell nuclear antigen from Pyrococcus furiosus
The polymerase subunit of DNA polymerase III of Escherichia coli. II. Purification of the alpha subunit, devoid of nuclease activities
Dynamics of DNA polymerase III holoenzyme of Escherichia coli in replication of a multiprimed template
The Escherichia coli preprimosome and DNA B helicase can form replication forks that move at the same rate
Characterization of a triple DNA polymerase replisome
Size classes of products synthesized processively by DNA polymerase III and DNA polymerase III holoenzyme of Escherichia coli
Single-molecule analysis reveals that the lagging strand increases replisome processivity but slows replication fork progression
Assembly of a chromosomal replication machine: two DNA polymerases, a clamp loader, and sliding clamps in one holoenzyme particle. I. Organization of the clamp loader
Clamp loader structure predicts the architecture of DNA polymerase III holoenzyme and RFC
Constitution of the twin polymerase of DNA polymerase III holoenzyme
Coupling of a replicative polymerase and helicase: a tau-DnaB interaction mediates rapid replication fork movement
Efficient in vitro replication of double-stranded DNA templates by a purified T4 bacteriophage replication system
PCNA connects DNA replication to epigenetic inheritance in yeast
Proliferating cell nuclear antigen (PCNA) is required for cell cycle-regulated silent chromatin on replicated and nonreplicated genes
AAA+: A class of chaperone-like ATPases associated with the assembly, operation, and disassembly of protein complexes
Structure at 2.8 A resolution of F1-ATPase from bovine heart mitochondria
Evolutionary relationships and structural mechanisms of AAA+ proteins
Mechanism of processivity clamp opening by the delta subunit wrench of the clamp loader complex of E. coli DNA polymerase III
Opening of the clamp: an intimate view of an ATP-driven biological machine
Mechanism of bacteriophage T4 DNA holoenzyme assembly: the 44/62 protein acts as a molecular motor
Fluorescence measurements on the E.coli DNA polymerase clamp loader: implications for conformational changes during ATP and clamp binding
The GTPase superfamily: a conserved switch for diverse cell functions
The internal workings of a DNA polymerase clamp-loading machine
Structural analyses of gp45 sliding clamp interactions during assembly of the bacteriophage T4 DNA polymerase holoenzyme. I. Conformational changes within the gp44/62-gp45-ATP complex during clamp loading
A model for Escherichia coli DNA polymerase III holoenzyme assembly at primer/template ends. DNA triggers a change in binding specificity of the gamma complex clamp loader
Structural and enzymatic studies of the T4 DNA replication system. II. ATPase properties of the polymerase accessory protein complex
ATP utilization by yeast replication factor C. I. ATP-mediated interaction with DNA and with proliferating cell nuclear antigen
Mapping the interaction of DNA with the Escherichia coli DNA polymerase clamp loader complex
Division of labor--sequential ATP hydrolysis drives assembly of a DNA polymerase sliding clamp around DNA
Crystal structure of the delta' subunit of the clamp-loader complex of E. coli DNA polymerase III
Ordered ATP hydrolysis in the gamma complex clamp loader AAA+ machine
The replication factor C clamp loader requires arginine finger sensors to drive DNA binding and proliferating cell nuclear antigen loading
Confirmation of the arginine-finger hypothesis for the GAP-stimulated GTP-hydrolysis reaction of Ras
Polymerase exchange during Okazaki fragment synthesis observed in living cells
Stoichiometry and architecture of active DNA replication machinery in Escherichia coli
Crystal structure of the chi:psi sub-assembly of the Escherichia coli DNA polymerase clamp-loader complex
Structure of the SSB-DNA polymerase III interface and its role in DNA replication
Devoted to the lagging strand-the subunit of DNA polymerase III holoenzyme contacts SSB to promote processive elongation and sliding clamp assembly
The chi psi subunits of DNA polymerase III holoenzyme bind to single-stranded DNA-binding protein (SSB) and facilitate replication of an SSB-coated template
A function for the psi subunit in loading the Escherichia coli DNA polymerase sliding clamp
The mechanism of ATP-dependent primer-template recognition by a clamp loader complex
Anatomy of a DNA replication fork revealed by reconstitution of SV40 DNA replication in vitro
Characterization of the five replication factor C genes of Saccharomyces cerevisiae
Replication factors required for SV40 DNA replication in vitro. I. DNA structure-specific recognition of a primer-template junction by eukaryotic DNA polymerases and their accessory proteins
Structural analysis of a eukaryotic sliding DNA clamp-clamp loader complex
How a DNA polymerase clamp loader opens a sliding clamp
Archaeal DNA replication: identifying the pieces to solve a puzzle
Open clamp structure in the clamp-loading complex visualized by electron microscopic image analysis
Distinct roles for ATP binding and hydrolysis at individual subunits of an archaeal clamp loader
Out-of-plane motions in open sliding clamps: molecular dynamics simulations of eukaryotic and archaeal proliferating cell nuclear antigen
Recognition of the ring-opened state of proliferating cell nuclear antigen by replication factor C promotes eukaryotic clamp-loading
The mechanical properties of PCNA: implications for the loading and function of a DNA sliding clamp
Structure of a small-molecule inhibitor of a DNA polymerase sliding clamp
Structure of the C-terminal region of p21(WAF1/CIP1) complexed with human PCNA
Structural and biochemical analysis of sliding clamp/ligand interactions suggest a competition between replicative and translesion DNA polymerases
The carboxyl terminus of the bacteriophage T4 DNA polymerase is required for holoenzyme complex formation
Structural analyses of gp45 sliding clamp interactions during assembly of the bacteriophage T4 DNA polymerase holoenzyme. III. The Gp43 DNA polymerase binds to the same face of the sliding clamp as the clamp loader
A molecular switch in a replication machine defined by an internal competition for protein rings
Interplay of clamp loader subunits in opening the beta sliding clamp of Escherichia coli DNA polymerase III holoenzyme
Structure and function of the bacteriophage T4 DNA polymerase holoenzyme
The 44P subunit of the T4 DNA polymerase accessory protein complex catalyzes ATP hydrolysis
A bipartite polymerase-processivity factor interaction: only the internal beta binding site of the alpha subunit is required for processive replication by the DNA polymerase III holoenzyme
A peptide switch regulates DNA polymerase processivity
A sliding-clamp toolbelt binds high- and low-fidelity DNA polymerases simultaneously
The dynamic processivity of the T4 DNA polymerase during replication
Coordination of multiple enzyme activities by a single PCNA in archaeal Okazaki fragment maturation
Creating a dynamic picture of the sliding clamp during T4 DNA polymerase holoenzyme assembly by using fluorescence resonance energy transfer
On the solution structure of the T4 sliding clamp (gp45)
Sliding clamp of the bacteriophage T4 polymerase has open and closed subunit interfaces in solution
Tracking sliding clamp opening and closing during bacteriophage T4 DNA polymerase holoenzyme assembly
Clamp loading, unloading and intrinsic stability of the PCNA, beta and gp45 sliding clamps of human, E. coli and T4 replicases
ATP binding to the Escherichia coli clamp loader powers opening of the ring-shaped clamp of DNA polymerase III holoenzyme
The Escherichia coli clamp loader can actively pry open the β-sliding clamp
Chl12 (Ctf18) forms a novel replication factor C-related complex and functions redundantly with Rad24 in the DNA replication checkpoint pathway
Identification of RFC(Ctf18p, Ctf8p, Dcc1p): an alternative RFC complex required for sister chromatid cohesion in S. cerevisiae
Replication protein A-directed unloading of PCNA by the Ctf18 cohesion establishment complex
Structure of a DNA:RNA hybrid duplex. Why RNase H does not cleave pure RNA
The structure of a ring-opened proliferating cell nuclear antigen-replication factor C complex revealed by fluorescence energy transfer
Mechanism of loading the Escherichia coli DNA polymerase III beta sliding clamp on DNA. Bona fide primer/templates preferentially trigger the gamma complex to hydrolyze ATP and load the clamp
Role of adenosine 5'-triphosphate hydrolysis in the assembly of the bacteriophage T4 DNA replication holoenzyme complex
ATP utilization by yeast replication factor C. II. Multiple stepwise ATP binding events are required to load proliferating cell nuclear antigen onto primed DNA
Bayesian shadows of molecular mechanisms cast in the light of evolution
The 'glutamate switch' provides a link between ATPase activity and ligand binding in AAA+ proteins
Multiple ATP binding is required to stabilize the "activated" (clamp open) clamp loader of the T4 DNA replication complex
Mechanism of proliferating cell nuclear antigen clamp opening by replication factor C
A central swivel point in the RFC clamp loader controls PCNA opening and loading on DNA
Only one ATP-binding DnaX subunit is required for initiation complex formation by the Escherichia coli DNA polymerase III holoenzyme
ATP utilization by yeast replication factor C. III. The ATP-binding domains of Rfc2, Rfc3, and Rfc4 are essential for DNA recognition and clamp loading
Dissecting the order of bacteriophage T4 DNA polymerase holoenzyme assembly
Examination of the role of the clamp-loader and ATP hydrolysis in the formation of the bacteriophage T4 polymerase holoenzyme
Dissection of the ATP-driven reaction cycle of the bacteriophage T4 DNA replication processivity clamp loading system
Bacteriophage T4 genetic homologies with bacteria and eucaryotes
Seeing the herpesvirus capsid at 8.5 A
Structural and functional similarities between the capsid proteins of bacteriophages T4 and HK97 point to a common ancestry
What does structure tell us about virus evolution?
Structural basis for the nuclease activity of a bacteriophage large terminase
Structure and inhibition of herpesvirus DNA packaging terminase nuclease domain
Evolutionary genomics of nucleo-cytoplasmic large DNA viruses
Lateral gene transfer, lineage-specific gene expansion and the evolution of Nucleo Cytoplasmic Large DNA viruses
Crystal structure of the processivity clamp loader gamma (gamma) complex of E. coli DNA polymerase III
Structure and mechanism of DNA polymerases
The structure of T. aquaticus DNA polymerase III is distinct from eukaryotic replicative DNA polymerases
Crystal structure of the catalytic alpha subunit of E. coli replicative DNA polymerase III
Structural analysis of bacteriophage T4 DNA replication: a review in the Virology Journal series on bacteriophage T4 and its relatives
dnaG gene product, a rifampicin-resistant RNA polymerase, initiates the conversion of a single-stranded coliphage DNA to its duplex replicative form
DNA or RNA priming of bacteriophage G4 DNA synthesis by Escherichia coli dnaG protein
Primase, the dnaG protein of Escherichia coli. An enzyme which starts DNA chains
Organization and evolution of bacterial and bacteriophage primase-helicase systems
A TOPRIM domain in the crystal structure of the catalytic core of Escherichia coli primase confirms a structural link to DNA topoisomerases
Structure of the RNA polymerase domain of E. coli primase
A DNA primase activity associated with DNA polymerase alpha from Drosophila melanogaster embryos
The single minichromosome maintenance protein of Methanobacterium thermoautotrophicum DeltaH contains DNA helicase activity
Isolation of the Cdc45/Mcm2-7/GINS (CMG) complex, a candidate for the eukaryotic DNA replication fork helicase
The Mcm2-7 complex has in vitro helicase activity
Activation of the MCM2-7 helicase by association with Cdc45 and GINS proteins
A new membrane-associated DNA replication protein, the gene 69 product of bacteriophage T4, shares a patch of homology with the Escherichia coli dnaA protein
Structural analysis of the inactive state of the Escherichia coli DNA polymerase clamp-loader complex


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About Time, not exactly a Rom Com
What up, it's Zac and I went to see "About Time" this weekend. I know, here's my man card. I was thinking it'd be a romantic comedy (ROM COM) with Rachel McAdams but I was actually pleasantly surprised. It has alot more depth and meaning and it's something that you could take your folks to see if they visit you for Turkey Day. I don't know if you should take your kids (they might have questions here and there) but it's a solid movie and Bill Nighy's wit as the Patriarch in this eccentric English family is second to none. It has its funny parts, it's sad parts and you walk away with an appreciation for those close to you and the road that you've taken versus the road you didn't take..... Go see "About Time", it's unique and well done and guys won't lose their man card for seeing it :)

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Who is Colm Feore playing in TASM 2? Sinister Six?
Is he simply Norman Osborn? Is he the Vulture and we're moving towards a Sinister Six film?
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With the recent casting of Colm Feore in the Amazing Spider-man 2, I can't stop wondering who he's playing. The popular theory seems to be Norman Osborn, which makes sense but I've also seen suggestions that maybe he's going to play the Vulture or even J. Jonah Jameson. I just wanted to outline my thoughts and see what you guys had to said so here goes.
Let's start with Norman Osborn. This seems to be the general consensus but I'm not so sure and hope it's not true. Now with Dane Dehaan cast as Harry Osborn, it would make sense that Norman should show up in this film but I think the series could benefit from him remaining in the shadows for another installment. I hope to see the Green Goblin kill Gwen in the third movie but don't think the character needs to be a major focus this time. I do think however he should be the Big Bad behind the scenes pulling the strings so this very well could be the role Feore was cast for but the idea I have could point elsewhere.
I'll call this the Sinister Six theory and I know it sounds overly ambitious but if we can have six heroes share a film, why not six villains? A Sinister Six could be tied together in the film through Oscorp's various research technolgies. With Jamie Foxx as Electro and Paul Giamatti as Rhino, Colm Feore could join their ranks as the Vulture. This makes three but also what about the man in the shadows played by Michael Massee last movie? Some had speculated that he was Norman Osborn but it seemed as if he teleported in and out of Connor's cell, maybe simply a trick of the eye and maybe he's Mysterio? Quite a leap I know but no more so than he has to be Norman Osborn. To round out the team, I suppose you'd have the Lizard return and either have Norman Osborn as the sixth member pulling the strings or add someone like the Scorpion as another villain easily explained by Oscorp tech. All these villains would battle Peter and in defeating them, he would figure out that Oscorp is behind it all, realizing the father of his "old friend" (so says the synopsis) is his greatest enemy. Whoever is cast as Norman could be shown at the end vowing that he need to take care of Spider-man himself leading to the Green Goblin in part 3.
I've also heard people kicking around the idea that Feore is going to play J. Jonah Jameson. Although I feel Colm Feore is a great actor, I really hope this isn't the role he was cast for. I just can't imagine him with a flattop and moustache chomping a cigar looking anything but ridiculous. J.K. Simmons did a wonderful job in the Raimi films and I know this series wants to distance itself from those but this is the one place it might be worth reconsidering.
Clearly you can tell where my hopes lie but I'm more likely totally wrong. Let me know what you guys think.
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Fashion appointments at PHOENIX magazine
By Amy Wilson
20th June 2014
Category: Consumer
PHOENIX magazine has made two appointments within its editorial team.
Nicole Smallwood has been appointed as fashion editor and Deborah LaTouche as menswear fashion editor. Both will be key figures in the continued development of the PHOENIX brand, creating conceptual, cinematic high-end fashion shoots.


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Peace of Mind.
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Mick Malaney President & Co-Founder
Sandy Malaney CEO & Co-Founder
Why Choose Big Hairy Dog?
Big Hairy Dog Information Systems offers services to customers that make us the most appealing choice for client support. We're available 24/7 with after-hours support. Our hours of operation are daily M-F from 6 AM to 6 PM, Saturday from 9 AM to 5 PM, and Sunday from 9 AM to 2 PM PST. After-hours support is available at any hour of the day or night. We do not believe in voicemail at BHD. Call us during regular business hours and you will reach a live person to assist you.
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Global software solutions provider Sapiens International, has significantly increased its investment in its North American insurance and retirement services practice.
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Pianist in Houston, Studio Musician. Ranked #1 pianist by GigMasters.***** ***** 2008 & 2009 GigMasters Rising Star Award ***** ***** ***** and highest feedback rating on this site.Quotes from customers:1. JP has an extraordinary talent first and foremost as an accomplished musician. From classical to smooth jazz and contemporary pop he contributed significantly to a wildly successful New Years...
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Data Science
Seeloz uses a cloud-based data mining engine to process billions of data signals from the typical data sources of any enterprise.
This data is the foundation of the entire solution. With it come new levels of business insights with respect to how the different components of the business are all correlated. With these correlations, we can accurately forecast performance and optimize it.
And what makes it better is that the predictive modeling and forecasting capabilities get stronger and more accurate as more data is collected through promotional campaigns and shopper interaction.
More Data Science = Better ROI. And nobody does Data Science like Seeloz.
Seamless Integration
With all these capabilities, you might expect all sorts of new tools, and enormous IT integration projects.
Not the case.
With Seeloz' non-disruptive approach, your existing business processes get empowered… not overhauled. Through our seamless IT integration process, Seeloz gets your enterprise up-and-running in days to weeks and you immediately start to see the tangible ROI.
Multi-Access-Level Consumption
Seeloz empowers every player in the organization with real-time dashboards, KPIs, and recommended actions.
Every player gets exactly what he needs based on her job responsibilities and visibility ranges, nothing more, nothing less.


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Student Health Insurance is located in the Health Services building, Room 201.
Office hours are Monday - Friday, 8:00 a.m. - 4:30 p.m. Summer hours may vary.
Mailing Address: Northern Illinois University Student Health Insurance Office Health Services Building, Room 101 DeKalb, IL 60115
Phone: 815-753-0122 Fax: 815-753-0965 Email: firstname.lastname@example.com
For Blue Cross Blue Shield questions, please call 855-267-0214.
DeKalb, Illinois 60115 | Regional Sites | Contact Us Emergency Information | Employment | Maps | Accessibility
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How to Change The Engine Oil on a Honda Rubicon?
The Honda Rubicon is a rugged ATV for riding everywhere and anywhere. However, riding in adverse conditions, such as in the rain, or on uneven surfaces, like mud and gravel, will require more freq...
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How to Change the Oil on the 2005 Honda Rubicon?
The 2005 Honda Rubicon is an ATV, or all-terrain vehicle. This type of vehicles can run on paved surfaces such as roads, or through muddy fields and forest trails. One of the key elements of keepin...
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Honda Rubicon Maintenance?
Preventative maintenance on your Honda Rubicon can help avoid having a good day quickly turn into a bad day. By regularly maintaining you ATV you can spend more time riding and less time with poten...
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How to Check the Oil in a 2001 Honda 500 Rubicon?
Engine oil is necessary for any gasoline-powered engine. Oil lubricates the many moving parts within the engine, keeping it from overheating due to friction. A low oil level can cause major engine ...
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2007 Honda Rubicon Information?
The 2007 FourTrax Foreman Rubicon is a utility-style all-terrain vehicle manufactured by Honda. The 2007 Rubicon originally sold for $7,349 and came with a six-month limited warranty and was a carr...
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Big Bend woman injured in crash
A 21-year-old Big Bend woman was injured when her vehicle collided with a dump truck in Vernon Wednesday afternoon, the Waukesha County Sheriff's Department said.
The woman was ejected during the crash and was taken by medical helicopter to Froedtert Hospital in Wauwatosa. She suffered serious head, facial and leg injuries, the sheriff's department said.
The crash occurred about 3:40 p.m. Wednesday.
The woman was southbound on Big Bend Drive and was turning onto eastbound Guthrie Drive when her vehicle collided with the dump truck, which was northbound on Big Bend Drive, the sheriff's department said.
The dump truck then struck a third vehicle, which was occupied by a Salem woman, 37, and her 11-year-old daughter, according to a sheriff's department news release.
The dump truck driver, a Waukesha man, 39, and the Salem woman and her daughter had minor injuries but were not taken to a hospital.


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Staffing Solutions Enterprises Expands Financial Staffing Services
Dale Kime Joins the Team as Business Development Representative
Dale Kime joins Ann Richardson, Branch Manager of Staffing Solutions, to help further expand and fulfill clients' needs in accounting and financial staffing and placement services. He brings over 13 years of business development experience in financial and accounting staffing services.
The new hire comes at a time of strategic expansion for Staffing Solutions Enterprises. Over the past several months, the company has grown to offer higher-level experienced candidates, particularly in accounting and financial roles.
"Dale's industry knowledge and experience, coupled with Ann's numerous years of client account management and recruitment at SSE, will allow us to expand our services to existing clients and branch out to deliver more comprehensive accounting services in the marketplace. We're excited to have him onboard," says SueAnn Naso, President of Staffing Solutions Enterprises.
Dale Kime will be responsible for developing relationships with existing clients in accounting and finance, and driving business development with new clients.
Dale obtained his Bachelor's Degree in Business Administration, with an emphasis in Finance from Cedarville University. He has been a member of the Institute of Management Accountants Cleveland East Chapter, where he was a past Board Member. He was also a member of the Greater Cleveland National Association of Credit Management.
About Staffing Solutions Enterprises:
We have a team of recruiting and HR experts who conduct thorough assessments and behavioral based screenings to match candidates and client workplaces. Whether our clients are looking for a permanent addition, a long-term contractor, or a short-term fill in, our focus is on making that perfect match because we understand clients want to add a team member-- someone who wants to be part of the company culture and create a WOW experience within their organization.
Jessica Morris
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By ELA RAE - Channel your inner gypset goddess with this stone embellished hand chain. 14k gold plating is accented with pyrite stone links and stunning encased moonstones. Stones elegantly lay on top of hand and are designed to accentuate the natural beauty of the wearer.
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text: Plot: Andrew is a troubled but creative teen with a keen visual eye and a high-quality HD camera. He is introverted and socially awkward. Along with his friends, Matt and Steve, they make a discovery that leads them all to acquire powerful telekinetic abilities. They're now capable of, well, almost anything. But things take a dark turn. Genre: Drama | Sci-Fi | Thriller IMDB rating: 7.4/10 from 28,064 users Directed by: Josh Trank Starring: Dane DeHaan, Alex Russell and Michael B. Jordan Release Name: Chronicle.2012.Directors.Cut.BluRay.1080p.DTS.x264-CHD Size: 8.69 GiB Video: x264 | 1920x1036 | 12420 Kbps Audio: English | DTS | 1536 Kbps Runtime: 88 mins Information: Code: http://www.imdb.com/title/tt1706593/ Download: Code: Extabit (1 GiB Parts): http://extabit.com/file/28dqc7xbrtw31/chd-chr0nicle720.part09.rar http://extabit.com/file/28dqc7xbrubrh/chd-chr0nicle720.part01.rar http://extabit.com/file/28dqc7xbsh2fx/chd-chr0nicle720.part03.rar http://extabit.com/file/28dqc7xbsh2fh/chd-chr0nicle720.part08.rar http://extabit.com/file/28dqc7xbsh2i5/chd-chr0nicle720.part04.rar http://extabit.com/file/28dqc7xbsh2hp/chd-chr0nicle720.part02.rar http://extabit.com/file/28dqc7xbsh2ct/chd-chr0nicle720.part05.rar http://extabit.com/file/28dqc7xbsh2cd/chd-chr0nicle720.part06.rar http://extabit.com/file/28dqc7xbsh2bx/chd-chr0nicle720.part07.rar Enjoy!


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An Australian child psychiatrist is facing child pornography charges after allegedly videotaping a child in a movie theatre washroom.
An angry father confronted a man in West Edmonton Mall on Sunday afternoon after he allegedly saw him recording a video of his son at a urinal in the Scotiabank Theatre's washroom.
The confrontation was captured in a video posted to Facebook.
"I was in the bathroom and this man was videotaping my son."
In the video, a man is surrounded by three others who push him to show them his cell phone.
"I was in the bathroom and this man was videotaping my son," says one of the men.
"Did you know recording in a public restroom is illegal?" adds another.
According to one bystander, the father eventually grabbed the man's phone and handed it to his partner.
"The mother goes 'I'm still looking, I'm still looking' and then after a couple of seconds, you kind of hear her let out this scream and that's when she said that she found the video of her son going to the bathroom. … I won't be able to forget that," witness Ashley Jackson told Global News.
Suspect was charged
One Facebook commenter who witnessed the exchange said she didn't know what happened at the time, but heard the boy's parents sounding upset.
"Mall security had the suspect fully surrounded waiting for the police and dad was making sure they weren't letting him go anywhere before the police arrived... kudos to the dad for keeping a level head during such an awful situation," she wrote.
Aaron Voon, 41, has been charged with possession of child pornography, making child pornography and voyeurism according to CBC News.
CORRECTION: An earlier version of this story indicated that Aaron Voon was an Edmonton resident. He is actually from Australia.
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About 73 percent of car seats are not installed correctly, but we can show you how to make sure your kid's ride is safe.
Every week, a child dies from a television or furniture tip-over. Check your house for furniture a child might try to climb and for TVs that could topple.
When a young child dies from a residential fire, a smoke alarm is usually either not working or not present. And carbon monoxide is not detectable, so your family needs both types of alarms.
Every 8 minutes, a child goes to the ER for medicine poisoning, and almost 9 times out of 10, the medicine belongs to an adult. Look at your house from a child's perspective to see what they can see or reach.
They come in musical greeting cards, remote controls, and key fobs. And of course, little kids pop them into their mouths in an instant. This happens more often than you might think-- there are about 10 phone calls a day to poison control centers.
To keep kids staying active and enjoying sports, young athletes and coaches need to know how to stay hydrated, stretch thoroughly, use the right equipment, and recognize the signs and symptoms of concussions.
Distraction impacts driving, walking, bath time, just about everything. Here's our challenge: take a month to notice times when you or your kids are distracted. Help each other remember to put down the phone (or whatever is diverting you) and focus on looking out for each other.


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ED Magedson – Founder
Ripoff Report
sanford brown institute islin nj675 us rt 1Islelin nj 08830 Internet, New Jersey United States of America
Does your business have a bad reputation?
Fix it the right way.
Corporate Advocacy Program™
I am looking for people who are just as angry as I am. This school is a scam they tell you misinformaton about courses and salaries. We need to do somehting about this as we are all in debt now and probably we never get a job making more then 12.00 an hour as this cost us $14,000.00. Please someone contact me if you feel the same way.
This report was posted on Ripoff Report on 01/22/2011 09:16 AM and is a permanent record located here: http://www.ripoffreport.com/reports/sanford-brown-institute-islin-nj/internet-new-jersey-08830/sanford-brown-institute-islin-new-jersey-this-school-is-a-scam-they-told-me-i-would-be-ma-685486. The posting time indicated is Arizona local time. Arizona does not observe daylight savings so the post time may be Mountain or Pacific depending on the time of year.
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Get the Wednesday newsletter
16:52 19th February 2011
Sheffield Wednesday exited the FA Cup on Saturday, going down 3-0 to Premiership outfit Birmingham City at St Andrew's.Goals from Jean Beausejour, Obafemi Martins and David Murphy did the damage, although the Owls did have moments of their own going forward.But the Blues had the clinical edge in the final third to progress to round six, leaving Wednesday to concentrate their efforts on climbing the League 1 table.After a solid start to the cup clash, the visitors saw that promise undone by Brum on six minutes, when Lee Bowyer burst down the left side of the penalty area and cut back for Beausejour to drill past Nicky Weaver.The Owls responded well, though, and were almost level inside five minutes but Neil Mellor's low effort was deflected wide for a corner.Play soon reverted to the other end and Keith Fahey forced a fine save from Weaver with a free kick right on the edge of the box.Weaver was called into action from the next attack, diving low to avert Beausejour's low drive round the post.Blues enjoyed most of the possession and the second goal that was looming duly arrived on 16 minutes. Martins showed the Owls' defence a clean pair of heels and advanced to slot home in front of the 4,000+ travelling fans from Sheffield.Birmingham suffered a blow when Alexander Hleb was stretchered off after a mistimed tackle by James O'Connor, before the Owls made a change when Richard Hinds came on for Mark Reynolds.Cameron Jerome went close for the home side as half time approached but Wednesday also threatened when Michael Morrison's shot from ten yards was blocked by Stephen Carrr.The Owls came closer still just before the break. Daniel Jones did well to shake off the attention of Stephen Carr and slide a low cross across the danger zone but the ball just evaded the loitering Mellor.Wednesday made two more changes for the second half, with Michael Morrison and O'Connor making way for Mark Beevers and Liam Palmer.But the pendulum stayed the way of Birmingham, who extended their lead on 52 minutes. A curling corner from the right fell kindly to Murphy to hit high into the net from close range and effectively put Birmingham out of reach.Martins could and should have made it four moments later but clipped over the bar following good work from substitute David Bentley.Wednesday rallied and enjoyed pockets of possession as the remainder of the game unfolded but couldn't breach the Blues' backline and the contest petered out with Birmingham's route to the quarter finals safely secure.Birmingham: Doyle, Murphy, Johnson (Mutch 63), Carr, Parnaby, Hleb (Bentley 26), Beausejour, Fahey, Bowyer, Jerome (Phillips 46), Martins
Unused substitutes: Foster, Larsson, Gardner, Zigic
Owls: Weaver, Spurr, R Johnson, M Morrison (Beevers 46), Reynolds (Hinds 28), Jones, Osbourne, O'Connor (Palmer 46), Teale, Morrison, Mellor
Unused substitutes: Jameson, Sedgwick, Potter, Heffernan


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ED Magedson – Founder
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RedboxBeach Blvd. Jacksonville, Florida United States of America
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Fix it the right way.
Corporate Advocacy Program™
This report was posted on Ripoff Report on 08/05/2012 02:28 PM and is a permanent record located here: http://www.ripoffreport.com/reports/redbox/jacksonville-florida-32224/redbox-redbox-redbox-charged-my-bank-account-6292-for-renting-3-movies-1-of-which-didn-922833. The posting time indicated is Arizona local time. Arizona does not observe daylight savings so the post time may be Mountain or Pacific depending on the time of year.
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Specifications can be changed without notice. Images are for illustrative purposes only. Retail Price (MSRP) means the manufacturer's/supplier's suggested retail price.
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- Meet with survivors at the forensic center and stay with the survivor throughout the evidentiary exam as desired by the survivor.
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Report: Krejci done with broken wrist
|05.05.10 at 11:23 pm ET
PHILADELPHIA — David Krejci was taken to a local hospital for treatment of an undisclosed injury following a hit by Philadelphia's Mike Richards in the first period. Krejci was on just his second shift of the game when he was drilled in the shoulder by Richards in the neutral zone. Just before the hit, Krejci found Milan Lucic, who then fed Miroslav Satan for the go-ahead goal at 5:45 of the first period.
"It was a clean hit and there are no issues there," Bruins coach Claude Julien said afterward.
The Bruins have not confirmed a CSNPhilly report that Krejci suffered a broken wrist and is scheduled to have corrective surgery in Baltimore.
The Bruins also played most of the game without defenseman Adam McQuaid, who was being evaluated by Bruins medical staff after the game for an undisclosed injury. Both Krejci and McQuaid played just two shifts before leaving and not returning.


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Scottish – Irish – Celtic
Celebrations honoring Celtic roots as well as customs have long been preferred. From the stirring cry of the bag pipe to the dress of formal tartan and kilt, weddings based upon elements of Celtic custom can provide long lasting memories. Aspects of these weddings could include part or all of the following:
Anam Cara - Literally, the Hearts other Half, this custom is a one-of-a-kind ceremony celebrating the development of a timeless love forever more. This event of the elements; to the Celts the four elements Earth, Fire, Water and Air were the structures on which a successful relationship were constructed.
Anam Cara
This tradition was adjusted as Christianity moved into the Celtic lands and also continues to be a remarkable way to include family and friends in your ceremony.
Handfasting - A ceremony dating back right to classical times, Handfasting is a custom of marriage prior to the availability of rings and rare-earth elements. The couple would take a piece of fabric or rope and before their friends and families, declare their love and intention by reciting a few words as well as binding themselves together symbolically with the cordage. It is from this practice we still refer to marriage as "tying the knot".
Oathing Stone -- What better area to place the guarantees of a lifetime as well as eternity than in the heart of a stone? The oathing stone is held by the groom and bride while their vows are stated, then in some traditions is tossed into a deep body of water to hold those promises for evermore. Today that stone could be maintained as a remembrance of this wedding.
Oathing Stone
The Quaich - Originally crafted from timber the twin handled Quaich was a Scotsman's canteen, mess kit and also drinking cup rolled right into one. King James of Scotland gave his betrothed Anne of Denmark a Quaich as a sign of his love for her during the marriage ceremony, from that point on the Quaich has actually been know as the "loving cup". This gorgeous event conveys the true blessings of Kith and also Kin to the couple.
The Quaich
Pinning of the Tartan
Pinning of the Tartan - A new bride is formally settled for right into the bridegroom's family through this event. Generally the oldest woman member of the groom's family offers a swatch of the family member 's Tartan to the new bride representing she is currently interwoven right into all the doings of the clan.
The presentation of the family sword - The martial origins of the Celtic individuals revolved around the protection of fireplace as well as home. This event is the acknowledgement of the male participants of the bride's household that they too currently have a brand-new partnership and also a new brother in arms.
Family Arms
At Life's Minutes weddings we can provide detailed support on Celtic/Scottish/Irish events, from construction of the ceremony to the last blessing in Gaelic we can assist you produce the day of your desires ...
Travel Costs
We Love to Travel! (however our accountant says we have to cover little things like gas, tires and maintenance), so to keep them from chewing our ears off our travel rates are as follows;
Albuquerque/Rio Rancho Metro Area (includes Corrales, Bernalillo, Isleta, Tijeras)
No Charge.
Fee per trip mileage is one way
30 – 60 miles
60—100 miles
101-175 Miles
Over 175 miles
Please call for details
Call 505-328-7397
Rev. Bruce Byers


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Alberta Main Street Alberta Main Street
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Alberta Main Street advances efforts to develop Alberta Street as a vibrant, creative & sustainable commercial district serving residents and visitors to our community.
By 2015, Alberta Main Street is a vibrant, funky and groovy organization that has made significant progress toward
- Creating a sustainable (social, economic, and environmental) business district.
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We succeeded in achieving this vision by integrating the following principles throughout our work:
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- Ensuring accessibility to all income levels.
- Prioritizing the safety of residents and visitors.
- Claire Gordon
- 503-683-3252
- Need admin access?
Would you recommend Alberta Main Street?


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Amid its disapproval by women groups, the government on Monday defended the decision to pass the anti-rape ordinance and urged 'everyone' to deal with the issue with 'utmost seriousness and sensitivity'.
"I appeal to everyone to deal with this issue with utmost seriousness and sensitivity," Union Finance Minister P Chidambaram said while addressing a press conference on Monday.
He negated claims that the recommendations of Justice Verma panel were rejected by the government.
"It is not correct to say that Justice Verma Committee report has not been accepted," Chidambaram said.
"There will be debate, consultation with political parties, which will give ample opportunities to reflect broadest perspective. The government wishes to assure everyone that further consultations will take place," he said.
"The government hopes stringent provisions in the ordinance will have a deterrent effect on potential criminals till a new law is enacted by Parliament," he said.
Speaking on the issue of marital rape, Chidambaram said: "Marital rape is a difficult issue on which we will have to hear the opinion of everybody."
President Pranab Mukherjee on Sunday signed the anti-rape law ordinance based on some of the recommendations of Justice Verma panel.
The ordinance, that came into effect as soon as Mukherjee signed it, will need to be passed by the Parliament within six months and is likely to be done in the Budget Session that begins on Feb 21.
Communist Party of India (Marxist) (CPI-M) leader Brinda Karat on Saturday said her party disapproved the new anti-rape law ordinance that was cleared by the Union Cabinet on Friday.
"We disapprove of the ordinance," Karat told media on Saturday.
"The content of the ordinance is doing injustice to the Justice Verma recommendations because it is highly selective in what it has included in the ordinance," she said.
"And in fact what it has included in some parts also is against what Justice Verma has recommended. We don't agree with this ordinance," Karat said.
Meanwhile, women right's groups also rejected the ordinance and had urged President Pranab Mukherjee not to sign it.
Human rights lawyer Vrinda Grover told media: "The ordinance is complete betrayal of the faith that people had put in the government."
"The President should not sign this ordinance," she said earlier.
The Union Cabinet on Friday cleared the new anti-rape law ordinance that mandates tougher jail terms for crimes against women, including the death penalty for extreme cases of sexual assault.
The ordinance incorporates some of the recommendations of the Justice Verma commission but ignores others such as cognizance of marital rape and amendments to the Armed Forces Special Powers Act (AFSPA).
Flaying the law enforcers and calling for police reforms, the Justice Verma Committee tasked by the government to suggest stringent laws against sexual violence submitted its "path-breaking" recommendations on Jan 23 to the Home Ministry, calling for enhancement of punishment to life sentence for the crime while stopping short of awarding death penalty.
The three-member committee headed by former Chief Justice of India J S Verma submitted its report to the government a month after it was formed following a national outrage over the brutal gang-rape of a 23-year-old medical student on a Delhi bus on December 16 and her subsequent death from the injuries.
Justice Leila Seth, former Chief Justice of Himachal Pradesh High Court, and former Solicitor General of India Gopal Subramanian, are the other two members of the committee.


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Home > Candy Recipes > Carob Truffles
Carob Truffles Recipe
Just plain healthy recipe...
Carob and artificial sweetener pair up with instant coffee and chopped peanuts to make very low sugar Carob Truffles.
Recipe Ingredients
4 cups carob - melted
1/8 teaspoon artificial sweetener
1/2 cup powdered instant coffee
1 cup finely chopped peanuts
Recipe Directions
Melt carob in double boiler. Add artificial sweetener to melted carob and stir well. You may use molds for fancy-shaped truffles or place mixture in a bowl to cool.
When cool enough to form a ball, form into 1-inch diameter balls and roll in either instant coffee or chopped peanuts (not both).
Place on waxed paper. Place truffles in refrigerator for 20 minutes to set.
Nutrition Nibbles
In addition to not having the negative effects of chocolate, carob is very nutritious. Carob contains as much Vitamin B1 as asparagus or strawberries; as much niacin as lima beans, lentils, or peas; and more Vitamin A than eggplant, asparagus, and beets. It also contains Vitamin B2, calcium, magnesium, potassium, and the trace minerals iron, manganese, chromium, copper, and nickel. It contains approximately 8 percent protein and is a good source of fiber. Compared to chocolate, carob is three times richer in calcium, has one third less calories and seventeen times less fat.
Nutrition Information
Serving size:
Calories: 57.2; Protein: 1.88g; Sodium: .097mg; Cholesterol: .806mg; Fat: 3.02g; Carbohydrates: 5.88g
Exchanges: 1/4 Bread/Starch; 1/2 Fat