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Structural basis of activity and allosteric control of diguanylate cyclase
Edited by Johann Deisenhofer, University of Texas Southwestern Medical Center, Dallas, TX, and approved October 19, 2004 (received for review August 19, 2004)
Recent discoveries suggest that a novel second messenger, bis-(3′→5′)-cyclic di-GMP (c-diGMP), is extensively used by bacteria to control multicellular behavior. Condensation of two GTP to the dinucleotide is catalyzed by the widely distributed diguanylate cyclase (DGC or GGDEF) domain that occurs in various combinations with sensory and/or regulatory modules. The crystal structure of the unorthodox response regulator PleD from Caulobacter crescentus, which consists of two CheY-like receiver domains and a DGC domain, has been solved in complex with the product c-diGMP. PleD forms a dimer with the CheY-like domains (the stem) mediating weak monomer–monomer interactions. The fold of the DGC domain is similar to adenylate cyclase, but the nucleotide-binding mode is substantially different. The guanine base is H-bonded to Asn-335 and Asp-344, whereas the ribosyl and α-phosphate moieties extend over the β2-β3-hairpin that carries the GGEEF signature motif. In the crystal, c-diGMP molecules are crosslinking active sites of adjacent dimers. It is inferred that, in solution, the two DGC domains of a dimer align in a two-fold symmetric way to catalyze c-diGMP synthesis. Two mutually intercalated c-diGMP molecules are found tightly bound at the stem–DGC interface. This allosteric site explains the observed noncompetitive product inhibition. We propose that product inhibition is due to domain immobilization and sets an upper limit for the concentration of this second messenger in the cell.
Cyclic nucleotides like cAMP or cGMP have been recognized as important low-molecular-weight signaling molecules. Although bacterial pathogens can interfere with cGMP signaling of their eukaryotic host cells (1), prokaryotes in general do not seem to use cGMP for signaling. In contrast, the cyclic dinucleotide bis-(3′→5′)-cyclic diGMP (c-diGMP) has been shown to regulate cell surface-associated traits and community behaviorlike biofilm formation in a number of bacterial species (2–5). The general importance of c-diGMP is underscored by the omnipresence of the diguanylate cyclase (DGC) domain (hitherto named GGDEF or DUF1) in bacterial genomes, where it occurs in various combinations with other sensory and/or regulatory modules (6, 7). Despite the wide distribution and obvious regulatory relevance of DGC proteins, structural and functional information about this class of regulators is largely missing.
In Caulobacter crescentus, pole remodeling during development is regulated by several polar sensor histidine kinases, which control the DGC activity of the response regulator PleD (8, 9). PleD is composed of a CheY-like receiver domain (D1), a CheY-like adapter domain (D2), and a DGC domain. Upon phosphorylation of D1, activated PleD sequesters to the differentiating pole, where it catalyzes the conversion of two molecules of GTP to c-diGMP (10). Thus, by coupling of activity and subcellular localization, the PleD readout is controlled in time and space. The conformational changes invoked by phosphorylation of response regulator receiver (RRR) domains have been studied thoroughly on CheY (11) and other single domain receiver proteins (for a review, see ref. 12). Little, however, is known about how these alterations are transmitted to the effector protein or domain, and only few intact multidomain response regulators have been analyzed structurally. Distinct mechanisms of activation, such as relief of active site obstruction [CheB (11, 13)], dimerization [DrrB, DrrD (14)], or both [NarL (15)], have been proposed.
Here, we report the structure of full-length PleD, which allows conclusions to be drawn regarding its mechanism of activation and product inhibition. Also, by providing a detailed view on the enzyme–product interactions, our study offers insight into the catalytic mechanism of this widespread family of regulatory proteins.
Materials and Methods
Overexpression and Purification. Full-length PleD with a C-terminal His-6 tag was overexpressed in Escherichia coli strain BL21 (DE3) pLysS. Purification was performed on a NiSO4-charged HiTrap chelating HP column (Amersham Pharmacia Biosciences) with elution at ≈200 mM imidazole. The pooled fractions were dialyzed overnight in 20 mM Tris·HCl (pH 8.0)/100 mM NaCl/1 mM DTT at 4°C. The protein solution was concentrated and, after clarification, loaded on a Superdex 200 HR 10/30 column(Amersham Pharmacia Biosciences), preequilibrated with the same buffer. PleD appeared as monomer, and fractions contributing to the peak were pooled. Selenomethionine-substituted PleD was expressed by using the metabolic inhibition pathway. The purification procedure was the same as for native PleD.
Enzymatic Assays. The assay for the measurement of the initial velocity of c-diGMP has been described (10). In short, GTP containing [α-32P]GTP (Amersham Pharmacia Biosciences; 5 nCi/microliter) was added to 50 μl of a solution containing 12.5 μg of PleD in 50 mM Tris·HCl (pH 7.8)/100 mM NaCl/10 mM MgCl2/5 mM 2-mercaptoethanol. Aliquots were taken at regular time intervals, and the product was assayed by thin-layer chromatography. For the product inhibition measurements, PleD was preincubated with c-diGMP at 25°C for 5 min. Care was taken to remove any bound c-diGMP from PleD carried over from the purification by extensive dialysis.
DGC activity was also assayed indirectly by monitoring the production of PPi by using an enzyme coupled spectrophotometric assay (16). The reaction mixture contained PleD in 20 mM Tris·HCl (pH 8.0)/100 mM NaCl/5 mM MgCl2/100 μM GTP and pyrophosphatase (500 milliunits/ml). The reaction was stopped by transferring a 100-μl reaction mixture into the phosphate assay reagent (pH ≈ 0; 1 ml final volume) containing molybdate and malachite green. The phosphate content in commercial GTP products had to be reduced from ≈5% to <0.5% by anion exchange chromatography to reduce background absorption (17).
Crystallography. Crystals were obtained at room temperature by hanging-drop vapor diffusion. For this, PleD at a nominal concentration of 200 μM = 10 mg/ml (assuming an ε280 of 9200 M–1·cm–1) in 20 mM Tris·HCl (pH 8.0)/100 mM NaCl/1 mM DTT/2 mM MgCl2/0.8 mM synthetic (10) c-diGMP was mixed with the reservoir (1.0 M glycine, pH 9.2/2% dioxane/14.5% polyethylene glycol 20 000) at a ratio of 1:1. Selenomethione-substituted crystals were obtained in the same manner, but by using a reservoir solution containing 1.0 M N-tris[hydroxymethyl]methyl-3-aminopropanesulfonic acid (TAPS) (pH 9.0), 2% dioxane, and 11% polyethylene glycol 20000. Diffraction data were collected from a single native (≈15 μm in diameter) and a single Se-Met substituted crystal (60 μm) at the Swiss Light Source, Paul Scherrer Institute, Villigen, Switzerland, and were processed with mosflm/scala (18). Eighteen selenium positions were identified by using shelxd (19). Phase refinement was performed by sharp/solomon (20) and was followed by two-fold averaging and phase extension by using dm (21). The model was built at the interactive graphics and refined by using refmac (18), imposing strict noncrystallographic symmetry constraints for the two copies in the asymmetric unit except for residues 117, 164, 168, and 404. The entire main chain is defined by electron density except for residues 137–146, 282–288, and the C-terminal His-tag.
Results and Discussion
Crystal Structure. The crystal structure of nonphosphorylated PleD has been solved by multiwavelength anomalous diffraction phasing on Se-Met substituted protein to 2.7 Å (Tables 1 and 2). The structure shows a linear arrangement of three structural domains (D1, D2, and DGC; Fig. 1a ) that are connected by single disordered loops. As anticipated, domains D1 and D2 show the typical RRR fold with (βα)5 topology. However, in both domains, the C-terminal helix α5 is considerably extended and protrudes from the globular domain, apparently enhancing the D1-D2 contact. Both domains closely resemble each other (rms deviation = 1.2 Å for 119 Cα). Quasiisologous contacts are formed between helix α5 of one domain and α4, β5, and α5 of the other domain. The C-terminal extensions of the α5 helices form apolar contacts, whereas their N-terminal parts are involved in several ionic interactions with α4 and β5 (Fig. 1b ).
Only D1 carries all of the residues required for RRR activation (11). These are primarily aspartates D9, D10, and the phosphoacceptor D53 of the "acidic pocket", but also residues K105, T83, and F102 that have been shown to undergo large structural changes upon activation (11). Strong density is bridging the three carboxylates of the acidic pocket and main-chain carbonyl 55. In line with its coordination geometry, this has been modeled as a Mg2+ ion. Comparison of D1 with other known RRR structures confirms that D1 is in the nonactivated conformation with the β4–α4 loop in an "outward" orientation. The structural changes accompanying activation in RRRs are known (22). In Fig. 1b , (nonactivated) PleD and activated FixJ (23) are superimposed. From this, one would predict that the largest conformational change upon PleD activation would occur in the β4–α4 loop with a concomitant shift (1–2 Å) in α4 that participates in the D1–D2 interaction. Another prominent change would be a rotameric switch of F102. Aromatic residues in this position have been shown to swing from a half-buried location into a pocket between α4 and the β-sheet that opens up on activation (11, 23). Notably and similar to CheB (24), F102 is located at a central position of the interface (Fig. 1b ). This argues for a repacking of the D1–D2 interface upon phosphorylation. How this may tie in with a global model of PleD activation is discussed below.
The DGC (GGDEF) domain consists of a five-stranded central β-sheet surrounded by helices (Fig. 1a ). Discounting two additional short β-hairpins (β0, β0′ and β3′, β3′′) and the N-terminal helix α0, the succession of secondary structure elements is (βααββαβαβ), for which the topology is identical and the arrangement closely similar to that of the catalytic core of adenylate cyclase (AC) (25) as well as to the "palm" domain of DNA polymerases (26) (Fig. 5, which is published as supporting information on the PNAS web site). Structural relatedness with AC has been proposed earlier by sequence threading (27). Obviously, the DGC domain is functionally related to AC and DNA polymerases in that it also catalyzes 3′–5′ phosphodiester formation. The GG(D/E)EF signature motif locates to the central β-hairpin (Fig. 1a ), which, as in AC and DNA polymerases, constitutes part of the active site.
In the crystal, PleD monomers are arranged to local dimers (Fig. 1c ). Isologous contacts are formed, with Y26 of D1 being tucked between helices α3 and α4 of D2 and the N terminus contacting α3. The small contact area (2 × 425 Å2) is consistent with the monomeric state of PleD in solution. Along the direction of their symmetry axis, dimers are associated head to head to form tetramers of 222 symmetry. Intriguingly, this interaction does not involve protein–protein contacts. Instead, as shown in Fig. 2a , c-diGMP is bound with its two guanine bases in a two-fold symmetric fashion to the active site formed by the adjacent monomers. This crosslinking of adjacent dimers represents a serendipitous crystallization artifact, because it provides comprehensive information about the product–enzyme interactions that appears relevant also for the situation in solution where the product would bridge the catalytic domains within a dimer (see below).
Active Site and Catalytic Mechanism. Fig. 2b shows that the guanine base is inserted in a pocket formed mainly by helices α1 and α2 of DGC. All of its polar atoms except O6 make specific interactions, explaining the observed GTP versus ATP discrimination (10). The N335 side chain forms H-bonds with N3 and N2, whereas N1 is recognized by the D344 carboxylate. Both residues are highly conserved among DGC domains. Furthermore, the base abuts the apolar side chains of L294, F331, and L347. The ribosyl and the α-phosphate are extending over the turn of the β2-β3-hairpin, with G369 being completely covered, explaining glycine conservation at this position. The 2′-hydroxyl is H-bonded to N335, and a phosphate nonester oxygen is in the vicinity of the main-chain amide 369.
On the basis of the observed complex structure, extrapolations regarding the binding mode of the GTP substrate to an individual DGC domain as well as the catalytic mechanism can be proposed (Fig. 2c ). Considering the specific interactions with the protein, it is most likely that the guanyl base does not change its position during catalysis. Furthermore, we assume that the ribosyl and α-phosphate moieties largely keep their positions. In this case, the β- and γ-phosphates of the substrate would be easily accommodated close to G368 of the β-hairpin and would probably be bound via a Mg2+ ion to E371, with R300 conferring further stabilization. For catalysis to occur, two substrate-loaded DGC domains have to arrange themselves in a similar way like the interdimer crosslink in the crystal (Fig. 2 b and c ). This might be facilitated by charge complementarity between D336 and R300 (Fig. 2b ). Subsequently, the 3′-OH group of the GTP substrate has to be deprotonated to allow an intermolecular nucleophilic attack onto the α-phosphate. E370 appears well poised for acting as a general base, whereas K332 would stabilize the charge of the developing pentacoordinated phosphoryl transition state and the pyrophosphate leaving group. All of the residues proposed to be of functional importance are highly conserved among DGC sequences.
Intriguingly, the proposed nucleotide-binding mode is different from that observed in AC (Fig. 2d ) and DNA polymerases. In both enzymes, the phosphate-binding site is provided by the short P-loop between β1 and α1. In AC, the base is lying on the β-hairpin and is recognized by residues of the second subunit of the heterodimer, whereas in DNA polymerases the base adopts even another position, which allows base pairing with the template strand (28). Is it conceivable that DGC can bind the substrate in a similar way as AC? This appears unlikely, because there would be no interactions with the base conferring specificity and contributing to binding affinity. An alternative model with the base bound to the DGC guanine-binding pocket and the terminal phosphates bound to the P-loop appears sterically unfeasible. Thus, there seems to be no reasonable alternative to the proposed model of the substrate–DGC complex, which nevertheless has to be confirmed experimentally. In AC, a magnesium ion coordinated by two aspartates (D396 and D440; Fig. 2d ) has been proposed to assist in the nucleophilic attack of the 3′-OH group onto the α-phosphate (25). Conspicuously, the DGC domain exhibits two invariant acidic residues (D327, E370) at very similar positions, but there is no indication for a bound metal, which is in line with the disorder of E370. In fact, a magnesium ion coordinated by these residues would be too distant from the substrate to be effective. E370 is proposed to act as a general base (see above), but the role of D327, if any, remains to be elucidated.
Allosteric Site and Product Inhibition. Unexpectedly, the crystal structure shows two product molecules bound to the D2/DGC interface (I-site; Figs. 1 and 3). The dimeric ligand is of compact roundish shape with mutually intercalated purine bases. Each of the two central guanyl moieties forms an intermolecular H-bond with a phosphate group. A similar structure has been observed for several crystal forms of c-diGMP (29, 30) and may be present also in solution. The ligand is bound to both D2 and DGC by a multitude of specific interactions. Hereby the common base–arginine pairing motif involving O6 and N7 of the guanyl (31) occurs three times (with R390, R359, and R178; Fig. 3b ). Tight binding is consistent with the observed copurification of c-diGMP during protein preparation. Intriguingly, kinetic data reveal strong product inhibition, with a K i (0.5 μM; Fig. 6, which is published as supporting information on the PNAS web site) about an order of magnitude lower than estimated for the cellular concentration of c-diGMP (32). Clearly, inhibition is noncompetitive (Fig. 6), i.e., independent of substrate concentration, and can thus be attributed to an allosteric effect of I-site binding.
Mechanistic Model of Regulation. PleD catalyzes a condensation reaction of two identical substrates (that are bound to two identical DGC domains) to yield a C2-symmetric product molecule. In solution, nonactivated PleD appears to be monomeric, as determined by analytical ultracentrifugation and gel filtration (not shown). For the enzyme to be efficient, dimerization appears necessary, because a reaction catalyzed by monomeric PleD would be severely limited by the macromolecular diffusion rate. Based on this reasoning, a simple mechanistic model of activation and product inhibition can be proposed (Fig. 4). Phosphorylation at D53 of D1 would induce repacking of the D1/D2 interface and reorientation of D1 with respect to D2. This in turn will enhance dimer formation mediated by isologous D1-D2 contacts across the interface. In fact, the dimeric structure of nonactivated PleD in the crystal may well resemble the active dimer, although with a suboptimal interface. This view is supported by the lack of activity of N-terminally His-tagged PleD (data not shown), which would be due to interference of the tag with the interface (Fig. 1c ). The low constitutive activity observed in vitro (10) would thus be due to a small fraction of nondissociated dimers. Consistent with this, the specific activity shows a strong dependence on the enzyme concentration (unpublished data).
The large distance between the substrate-binding sites of the crystal dimer (Fig. 1c ) would not permit catalysis. However, allowing for flexibility of the DGC domains relative to the stem (a realistic assumption considering the small D2/DGC interface area of 690 Å2), a "closed" dimer conformation competent for catalysis can be readily modeled (schematically shown in Fig. 4), in which a complete two-fold symmetric active site is formed as observed for the DGC/DGC′′ pair (Fig. 2b ). In the context of this activation-by-dimerization model, the observed allosteric product inhibition can easily be explained (Fig. 4). Stabilization of the D2/DGC interface by product binding to the I-site would simply prevent encounter of the substrate-binding sites (inhibition by domain immobilization).
The tight feedback regulation of PleD by product inhibition probably demonstrates the importance of imposing an upper limit on the concentration of the second messenger c-diGMP. It remains to be shown that, as predicted by the model, inhibition indeed constitutes an overriding principle, i.e., that inhibition is independent of the phosphorylation state of the enzyme. The structure of PleD provides insight into the molecular interactions with c-diGMP that may recur in the recognition of this ligand by other proteins. The quest for downstream targets of this messenger that appears to be ubiquitous in prokaryotes (6) has started.
We thank T. Bickle, Z. Markovic-Housley, O. Mayans, and C. Peneff for critical reading of the manuscript; J. Saito for crystallization optimization; and D.V. Palanivelu and the staff of beamline X06SA of the Swiss Light Source synchrotron (Villigen, Switzerland) for assistance with data acquisition. The work was supported by Swiss National Science Foundation Grant 31-59050.99 (to U.J.).
↵ § To whom correspondence should be addressed. E-mail: tilman.schirmer{at}unibas.ch.
Author contributions: B.G., U.J., and T.S. designed research; C.C., R.P., D.S., N.C.A., and T.S. performed research; D.S. and N.C.A. contributed new reagents/analytic tools; C.C., R.P., U.J., and T.S. analyzed data; and C.C., U.J., and T.S. wrote the paper.
This paper was submitted directly (Track II) to the PNAS office.
Abbreviations: c-diGMP, bis-(3′→5′)-cyclic di-GMP; DGC, diguanylate cyclase; RRR, response regulator receiver; AC, adenylate cyclase.
Data deposition: The atomic coordinates and structure factors have been deposited in the Protein Data Bank, www.pdb.org (PDB ID code 1W25).
- Copyright © 2004, The National Academy of Sciences
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These are definitely "not your mothers" golf clothes. Foray Golf, inspired by the new and now, was founded in 2016 by fashion industry veterans who love to golf. The modern designs provide a fresh twist on golf fashion. The limited edition fashion apparel is unexpected, fun, and performs both on and off the course.
When fashion and function collide, a truly magical piece of clothing is created. The Foray Golf Rockstud Layering Long Sleeved Top adds edge to your wardrobe on and off the course. This lightweight marl neoprene mid-layer features a Gold and Navy striped metallic rib hem and crew neckline.

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New and classic shoes
Open heel
The heel is comfortable
Material is velvet
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Pick two pairs at 3000
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Only available in this colors
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Available in this colors
The heel is not too high, very comfortable
Material is leather
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Make 1 ½-Pound Recipe with bread machines that use at least 3 cups flour, or make 2-Pound Recipe with bread machines that use 4 cups flour.
Measure carefully, placing all ingredients except raisins in bread machine pan in the order recommended by the manufacturer. Add raisins at the Raisin/Nut signal.
Select Whole Wheat or Basic/White cycle. Use Medium or Light crust color. Do not use delay cycles Remove baked bread from pan, and cool on wire rack.
1 Slice: 180 calories (20alories from fat) 2 fat (0g saturated) 0mg cholesterol 290sodium 38 carbohydrate (2g dietary fiber) 4g protein
Success Tip: We found from our testing that for good texture and volume, slightly less yeast is needed in the 2-pound loaf than is needed in the 1 1/2-pound loaf.
Success Tip: If your bread machine doesn't have a Raisin/Nut signal, add the raisins 5 to 10 minutes before the last kneading cycle ends. Check your bread machine's use-and-care book to find out how long the last cycle runs.
Dried cranberries used instead of the golden raisins would add a slight tart flavor to this bread. Or use regular dark raisins instead of the golden ones for the same sweet flavor and a nice color contrast.
From "Betty Crocker's Best Bread Machine Cookbook, The Goodness of Homemade Bread the Easy Way." Text Copyright 1999 General Mills, Inc. Used with permission of the publisher, Wiley Publishing, Inc. All Rights Reserved.
This Golden Raisin Bread recipe is from the Betty Crocker's Best Bread Machine Cookbook Cookbook. Download this Cookbook today.
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What doesn't kill her makes her stronger. Kelly Clarkson returned to The Voice Monday night just days after having surgery to remove her appendix. Clarkson revealed on Twitter the day after hosting the Billboard Music Awards last week, that she "broke down in tears after the show from pain," and that she "flew home directly after the event" to have the procedure. On Sunday, Clarkson updated her fans as she tweeted: "Recovering after surgery super duper sucks. Turns out I don't "rest" well (so bored), pain meds make me feel a different kind of horrible to where I question if I'd just rather feel the pain instead, & 2 out of the 3 scars feel as if Lord Voldemort is always near." Despite having a hard time recovering, Clarkson is carrying on like the pro that she is. Fans noticed that there was a lot less of Clarkson during Monday night's show and that she wasn't her usual self, but they were all very supportive. One person tweeted: "@kelly_clarkson don't ever stop. She just had surgery, and still on #TheVoice. And she performed two times last week and didn't miss a beat. She tough."
'The Real' co-host Jeannie Mai opens up about being a victim of sexual abuse: 'You never heal, you just continue to heal'
Danny Glover says reparations for slavery is a 'moral, democratic and economic imperative'
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Western Cape
The Western Cape of South Africa is among some of the most beautiful vineyard locations in the world - vivid green pastures tucked between granite and sandstone cliffs, sapphire-blue ocean, and the bright white edifices of 300 year old colonial homesteads. One of the larger geographical units within the South African Wine of Origin classification system, most of the nation's wines come from this region.
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Erasmus Family, Cape Wine Company, 2014
This wine is a blend of Carignan, Mourvèdre, Grenache and Shiraz. It is a typical Mediterranean blend reflecting the elegance of the 2014 vintage with ripe, soft tannins. Only 3000 bottles were produced, making it quite the exclusive blend.Learn More£39.50
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A Psalm of David. The Lord is my shepherd; I shall not want. He makes me lie down in green pastures. He leads me beside still waters. He restores my soul. He leads me in paths of righteousness for his name's sake. Even though I walk through the valley of the shadow of death, I will fear no evil, for you are with me; your rod and your staff, they comfort me. You prepare a table before me in the presence of my enemies; you anoint my head with oil; my cup overflows. ... Share Your Faith Products
My son, do not forget my teaching, but let your heart keep my commandments, for length of days and years of life and peace they will add to you. Let not steadfast love and faithfulness forsake you; bind them around your neck; write them on the tablet of your heart. So you will find favor and good success in the sight of God and man. Trust in the Lord with all your heart, and do not lean on your own understanding. ... Share Your Faith Products
"You shall make two cherubim of gold, make them of hammered work at the two ends of the mercy seat. "Make one cherub at one end and one cherub at the other end; you shall make the cherubim of one piece with the mercy seat at its two ends. "The cherubim shall have their wings spread upward, covering the mercy seat with their wings and facing one another; the faces of the cherubim are to be turned toward the mercy seat.read more. Bible Verse Wall Art
"You shall have no other gods before Me. "You shall not make for yourself an idol, or any likeness of what is in heaven above or on the earth beneath or in the water under the earth. "You shall not worship them or serve them; for I, the LORD your God, am a jealous God, visiting the iniquity of the fathers on the children, on the third and the fourth generations of those who hate Me,read more. Christian Gifts
and in the cutting of stones for settings, and in the carving of wood, that he may work in all kinds of craftsmanship. "And behold, I Myself have appointed with him Oholiab, the son of Ahisamach, of the tribe of Dan; and in the hearts of all who are skillful I have put skill, that they may make all that I have commanded you: the tent of meeting, and the ark of testimony, and the mercy seat upon it, and all the furniture of the tent, the table also and its utensils, and the pure gold lampstand with all its utensils, and the altar of incense, the altar of burnt offering also with all its utensils, and the laver and its stand,
It was carved with cherubim and palm trees; and a palm tree was between cherub and cherub, and every cherub had two faces, a man's face toward the palm tree on one side and a young lion's face toward the palm tree on the other side; they were carved on all the house all around. From the ground to above the entrance cherubim and palm trees were carved, as well as on the wall of the nave. The doorposts of the nave were square; as for the front of the sanctuary, the appearance of one doorpost was like that of the other. The altar was of wood, three cubits high and its length two cubits; its corners, its base and its sides were of wood And he said to me, "This is the table that is before the LORD." The nave and the sanctuary each had a double door. Each of the doors had two leaves, two swinging leaves; two leaves for one door and two leaves for the other. Also there were carved on them, on the doors of the nave, cherubim and palm trees like those carved on the walls; and there was a threshold of wood on the front of the porch outside. There were latticed windows and palm trees on one side and on the other, on the sides of the porch; thus were the side chambers of the house and the thresholds. Share Your Faith Products
For the last 5 years, the in-house design team at the Faithlife Corporation has illustrated one Bible verse every day. This art has found its way onto t-shirts, magnets, and postcards—and now, a beautiful picture book. In print for the first time, art from Faithlife's Verse of the Day series paired with uplifting devotionals will encourage and inspire you. Christian Gifts
What does the Bible say about creativity? The very first verse of Scripture actually describes a creative act as "God created the heavens and the earth." Additionally, from Genesis, the Bible says, "God created man in his own image." This could be interpreted as we are creators just as God created us. We create relationships, art, homes, cities, nations and many more things including inventions even yet to be known! The creative spirit is certainly favored by God and our fellow man. Learn more from the collection of Bible verses about creativity below! Bible Verse Wall Art
2 Timothy 3:16 tells us that ALL Scripture is inspired by God and useful for teaching, rebuking, correcting, and training in righteousness. The Bible is a work of God that was written by human men that were inspired by God. What better source of inspiration than our Creator! Whether you are looking for motivation, encouragement, reassurance, or peace, the Bible should be the first resource you turn to! Christian Gifts
The Lord said to Moses, "See, I have called by name Bezalel the son of Uri, son of Hur, of the tribe of Judah, and I have filled him with the Spirit of God, with ability and intelligence, with knowledge and all craftsmanship, to devise artistic designs, to work in gold, silver, and bronze, in cutting stones for setting, and in carving wood, to work in every craft. ...
Then God said, "Let us make man in our image, after our likeness. And let them have dominion over the fish of the sea and over the birds of the heavens and over the livestock and over all the earth and over every creeping thing that creeps on the earth." So God created man in his own image, in the image of God he created him; male and female he created them.
Solomon the son of David established himself in his kingdom, and the Lord his God was with him and made him exceedingly great. Solomon spoke to all Israel, to the commanders of thousands and of hundreds, to the judges, and to all the leaders in all Israel, the heads of fathers' houses. And Solomon, and all the assembly with him, went to the high place that was at Gibeon, for the tent of meeting of God, which Moses the servant of the Lord had made in the wilderness, was there. (But David had brought up the ark of God from Kiriath-jearim to the place that David had prepared for it, for he had pitched a tent for it in Jerusalem.) Moreover, the bronze altar that Bezalel the son of Uri, son of Hur, had made, was there before the tabernacle of the Lord. And Solomon and the assembly sought it out. ... Bible Verse Wall Art
What does the Bible say about creativity? The very first verse of Scripture actually describes a creative act as "God created the heavens and the earth." Additionally, from Genesis, the Bible says, "God created man in his own image." This could be interpreted as we are creators just as God created us. We create relationships, art, homes, cities, nations and many more things including inventions even yet to be known! The creative spirit is certainly favored by God and our fellow man. Learn more from the collection of Bible verses about creativity below! Christian Canvas Art
Since the advent of printing, the sale of reproductions of pious works has been a major element of popular Christian culture. In the 19th century, this included genre painters such as Mihály Munkácsy. The invention of color lithography led to broad circulation of holy cards. In the modern era, companies specializing in modern commercial Christian artists such as Thomas Blackshear and Thomas Kinkade, although widely regarded in the fine art world as kitsch,[4] have been very successful.
He also made two pillars for the front of the house, thirty-five cubits high, and the capital on the top of each was five cubits. He made chains in the inner sanctuary and placed them on the tops of the pillars; and he made one hundred pomegranates and placed them on the chains. He erected the pillars in front of the temple, one on the right and the other on the left, and named the one on the right Jachin and the one on the left Boaz. Bible Verse Wall Art


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Animal Paper Plates
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Washington Squirrel University: Week 3
NICK SANDIFER, Evergreen cartoonistMarch 9, 2018Filed under Community, Involvement, Life, Washington Squirrel University
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Man faces to serve nine months in jail following plea deal
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Our plumbers are licensed, bonded, and insured and meet our rigid trust certification standards. Our services are guaranteed in writing against defects in materials and workmanship. So if you need to repair, maintain, or improve your plumbing or septic system, you can rely on Parker & Sons. Our plumbing service includes a 100% money back guarantee and our customer care policies are unmatched in the industry.
Estimating a plumbing job is best left to the professionals. However, our guides linked to throughout this article are the best first step to understanding pricing. Understanding basic plumbing is an excellent second step. Not only does this help you diagnose potential problems before they become costly ones, but itll help you understand what a plumber does.
Mr. Rooter® Plumbing is the go-to plumber for homeowners across the country. Why? Our plumbing contractors provide courteous, friendly, affordable, and effective residential plumbing services. We take our mission to exceed our customers' expectations seriously! From on-time arrivals to licensed and certified plumbing experts, we make sure homeowners are getting the very best plumbing services. You can have total peace of mind and confidence when you call on us! We stand behind our work and go the extra mile to get the job done quickly so you can get back to enjoying your home.
"Don't assume that every Mr. Fix-it advertising his services in the local Pennysaver or on Craigslist is a licensed plumber," says one New York plumber. There is no national standard for issuing licenses. Some plumbers are licensed by the state, others by the counties they work in. Check with your local city hall or chamber of commerce. They should be able to direct you to the appropriate source for a list of licensed plumbers in your area.
Plumbing originated during ancient civilizations such as the Greek, Roman, Persian, Indian, and Chinese cities as they developed public baths and needed to provide potable water and wastewater removal, for larger numbers of people.[6] Standardized earthen plumbing pipes with broad flanges making use of asphalt for preventing leakages appeared in the urban settlements of the Indus Valley Civilization by 2700 BC.[7] The Romans used lead pipe inscriptions to prevent water theft. The word "plumber" dates from the Roman Empire.[8] The Latin for lead is plumbum. Roman roofs used lead in conduits and drain pipes[9] and some were also covered with lead. Lead was also used for piping and for making baths.[10]
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On his first visit to our home, Eduardo quickly fixed our plumbing clog and also asked several questions to diagnose WHY we might be having these recurrent backups. He's the rare plumber who does more than just fix the symptom! He's also very conscientious about cost, informing us through the process of any work that would impact our bill. He offered us choices, and explained thee cost and benefit of each so we can make informed decisions and understand potential outcome: He's a plumber AND a plumbing consultant!
Hii need you guys help! Once my thermostat drop below the set temp , my furnace does not turn on and I was wondering what's wrong with it. It happens mostly on cold nights. Sometimes I would have to go in the basement and turned the furnace power switch off then on and will come on once I switch it off then on.i also change the thermostat recently, so I don't think that's the problem
Water systems of ancient times relied on gravity for the supply of water, using pipes or channels usually made of clay, lead, bamboo, wood, or stone. Hollowed wooden logs wrapped in steel banding were used for plumbing pipes, particularly water mains. Logs were used for water distribution in England close to 500 years ago. US cities began using hollowed logs in the late 1700s through the 1800s. Today, most plumbing supply pipe is made out of steel, copper, and plastic; most waste (also known as "soil")[22] out of steel, copper, plastic, and cast iron.[22]
On average, we hire only 1 out of every 10 plumbers in Los Angeles who apply to join our plumbing services team. In other words, we don't hire amateurs or inexperienced plumbers. Because of the intensive training program, each plumber becomes proficient in carrying out plumbing services to the utmost satisfaction of our customers. Background checks are performed on all personnel, and all Mike Diamond employees are drug free and wear photo identification. So where do the 90% that we don't hire go? Somewhere else.
The word "plumber" dates from the Roman Empire.[3] The Latin for lead is plumbum. Roman roofs used lead in conduits and drain pipes[4] and some were also covered with lead, lead was also used for piping and for making baths.[5] In medieval times anyone who worked with lead was referred to as a plumber as can be seen from an extract of workmen fixing a roof in Westminster Palace and were referred to as plumbers "To Gilbert de Westminster, plumber, working about the roof of the pantry of the little hall, covering it with lead, and about various defects in the roof of the little hall".[6] Thus a person with expertise in working with lead was first known as a Plumbarius which was later shortened to plumber.
Plumbing is any system that conveys fluids for a wide range of applications. Plumbing uses pipes, valves, plumbing fixtures, tanks, and other apparatuses to convey fluids.[1] Heating and cooling (HVAC), waste removal, and potable water delivery are among the most common uses for plumbing, but it is not limited to these applications.[2] The word derives from the Latin for lead, plumbum, as the first effective pipes used in the Roman era were lead pipes.[3]
My parents don't speak much English, but he was very patient with us today while I translated for them, which I truly appreciate. When he came by today, he explained what he was doing step by step; he was very clear about the pros, cons, and costs associated with our different options. Eduardo's attitude is great, and I feel like he genuinely wanted to help us. Definitely the kind of person who you want to work with. My parents have already passed his information onto all of their church friends, and I have him saved as a contact in case I ever need help again. Thank you Eduardo! :)
"Kennith, saved me about $1,500 by steam cleaning my main sewer lateral (4" line from under house to the city sewers), versus other options for the same work, and about $6,000 if I had listened to the guys that wanted me to replace my lateral. Had ALL the equipment you could ever not even expect in the back of his truck. Serious specialty equipment. I was very impressed. So, CALL Ken Vasil 817-983-1348 in Fort Worth and surrounding area's & keep him well employed as an excellent specialty plumber!"
Apprentice plumber: Apprenticeship programs generally provide the most comprehensive training for novice plumbers. Either union locals and their affiliated companies or nonunion contractor organizations can administer this training. Apprenticeships typically consist of at least three to four years of paid, on-the-job training and some hours of related classroom instruction.
Cast iron and ductile iron pipe was long a lower-cost alternative to copper, before the advent of durable plastic materials but special non-conductive fittings must be used where transitions are to be made to other metallic pipes, except for terminal fittings, in order to avoid corrosion owing to electrochemical reactions between dissimilar metals (see galvanic cell).[18]
My plan was to have an old kitchen faucet head replaced with a new one I had bought, a 5 minute job at most. The plumber told me it would first be a $28 assessment fee and then a "job " cost but they could not quote phone costs or give me any indication of a range cost. I declined to use them and to find a handyman instead. Plumbers stories significantly overcharging seem prevalent and I could see it could well happen to me. I got a handyman who did the job in 3 minutes flat! The Plumbing Costs Chart you sent out is good, it shows it would probably have cost me $200 for 5 minutes plus $28 assessment fee.
JJ's Plumbing Repair has no control over, and assumes no responsibility for, the content, privacy policies, or practices of any third party web sites or services. You further acknowledge and agree that JJ's Plumbing Repair shall not be responsible or liable, directly or indirectly, for any damage or loss caused or alleged to be caused by or in connection with use of or reliance on any such content, goods or services available on or through any such web sites or services.
Our qualified, certified and experienced Denver plumbers specialize in sewer line cleaning, sewer video inspection, sewer and water taps, trenchless repair and pipe bursting. We also locate and handle leaks, and the repair and replacement of both sewer and water lines. Our customer-first service and quality of work earned us the Angie's List Super Service Award from 2012-2017, and an A-plus rating with the Better Business Bureau. We have the knowledge and drive to complete all your plumbing services.
Sump pumps are pivotal during heavy rainstorms and prevent many issues that would require plumbing repairs. For example, when it rains, groundwater around your house's foundation will be guided into a bordering drain system. From there, the water flow is directed into pipes and diverted to the sump pit, well, and sump pump for collection/elimination.
Finally I get to write my fantastic review of Eduardo after the holidays! I had problems with my toilet, the tank refilling every hour. In addition, i noticed one of the rubber pieces disintegrating at the base. I did not trust myself after googling how to fix this problem. After looking at the fantastic reviews on YELP for this company, I chose to call Eduardo. Eduardo was very nice and gave me a proposed estimate on the phone, also giving me options if I had wanted to purchase any pieces on my own to save cost. He came the day after and immediately got to work. Eduardo was neat, wearing shoe covers into the house, and also wiping down the tank after unscrewing it off the toilet. I really appreciated this. Eduardo also knew everything he was doing without hesitation. He was super friendly and gave me plenty of tips as he was working away! Later that evening there was an issue, of the tank water leaking into the toilet. After calling Edward, he came right back and replaced one of the pieces I had purchased with one of his preferred brands. I was shocked that a brand new piece could be defective. But Eduardo was quite certain that was the problem! Sure enough, after Eduardo reinstalled that piece, my toilet has been running like a charm! Eduardo is surely the person I will call for future plumbing needs! He really cares about his customers and the work he does for them! And btw, even though he had to come back a second time, he had the best of attitudes, and only wanted to get the job done right! Thanks, Eduardo!! Happy New Years!!!!! A definite A+++++ job!!! Lydia of Irvine


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The EPL carries on right now with 6 matches slated to generally be performed. Chelsea will likely be facing out with Tottenham although Crystal palace will probably be facing out with manchester united.We've finished Examination for yourself that…
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Pebble Beach Golf Links and the Pebble Beach Resort hold special reverence in the golf world. Every golfer who hasn't been wants — and ought — to make a pilgrimage there. With the exception of St. Andrews, no place in golf conjures the sport's allure like Pebble Beach. To utter the name is to summon images of water crashing the sea wall, land's end par threes like the 7th and 17th, a glorious fini
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The smart Trick of Profit Trading That Nobody is Talking About
For instance, the SHA-256 of the term BUTTERFLY (origin ) is 8c62ace4f9ef8ccd08ca6fb992a8524bb7dbdc0530654bd254c9da07a660949a (HASH). This seemingly random string of letters and numbers contains three important properties:
Bitcoin mining involves three factors: the cube, the mining issue and a random number. Heres how it all comes together:
Imagine our cube consists of the term BUTTERFLY discussed earlier. In reality, the block would contain a list of recent, unverified transactions, but lets keep it simple. In order for the block to be solved, bitcoin uses a simple test: If the HASH consequence of the block starts with a certain number of zeros, then the cube is considered verified.
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For our example, lets say that we have a mining difficulty of simply two, ie, our HASH must begin with two zeros. .
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The problem: BUTTERFLY will return the same HASH, and it doesnt begin with two zeros. So what we need is the third factor, a random number (called a NONCE). We carry this number, combine it with BUTTERFLY, and HASH again. If it doesnt start with two zeros, we change the number and try again, and since changing one little number changes the entire HASH result, there's absolutely no way to forecast the number well need to solve this! .
We repeat this procedure over and over until we find a number that, when combined with BUTTERFLY, gives us a HASH that starts with two zeros. That number is your solution to the block. Here are some tries:
This arduous procedure of randomly trying to find a number that gives the solution is the thing that creates bitcoin mining such a computationally expensive process, and as more miners join the network, the tougher it gets. As of November 2017, a regular home computer working alone, ie, not an application-specific integrated circuit (ASIC) and not a part of a cloud mining network, could require 2.7 million years to mine one block. .
This has caused the growth of ASIC computers constructed specifically for mining and also to an original site increase in cloud mining.
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CPU mining. In the first days of bitcoin, mining issue was reduced and not a great deal of miners were competing for cubes and rewards. This made it rewarding to utilize your computers own central processing unit (CPU) to mine bitcoin. However, that strategy was soon replaced by GPU mining.
GPU mining. A graphics processing unit (GPU) is a powerful processor whose sole objective is to assist your own computers graphics card in rendering 3D graphics. GPUs are not constructed for executive decisions (like CPUs) however to be somewhat good labourers, hence GPUs can execute over 800 times more instructions in the exact same amount of time as a CPU.
FPGA mining. Next navigate to this website came mining with field-programmable gate arrays (FPGAs). These greatly outperformed GPUs and CPUs in the mining process as FPGAs are processors which can be programmed to execute certain instructions and only those instructions (instead of being repurposed for mining, like right here GPUs were).
ASIC mining. Similar to FPGAs, application-specific integrated circuits are processors designed for a particular purpose, in our situation mining bitcoin, and nothing else. ASICs for bitcoin were introduced in 2013 and, as of November 2017, they're the best processors out there for mining bitcoin and they outperform FPGAs in power consumption. .
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Mining pools. To cancel the problem of mining a block, miners started organising in pools or cloud mining networks. Whenever a miner in one of these pools solves a block, the payoff is shared with everyone in the swimming pool in a ratio representative of just how much work you put into the pool (even though you personally never solved the puzzle). .
Cloud mining. Clouds provide prospective miners the capability to buy mining channels in a remote data centre location. There are many obvious advantages, the most obvious being: no electricity costs, no excess heat and nothing to sell when you decide to hang your virtual pickaxe.
Once miners receive bitcoin, they are given a virtual key to the bitcoin addresses. You can use this electronic key to access and confirm or approve transactions.
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Desktop pockets. Software like Bitcoin Core allows you to send and store bitcoin addresses and connects to the network to track transactions.
Online wallets. Bitcoin keys are stored online by exchange programs such as Coinbase or Circle and can be accessed from anywhere.
Mobile wallets. Programs like Blockchain store and encrypt your own bitcoin keys so that you can make payments using your cellular device.
Paper wallets. Some sites offer paper wallet solutions, generating a piece of paper using two QR codes on it. One code is your public address where you receive bitcoin and the other one is the personal address you can use for spending.


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Asbestos-Laced Makeup: Children's Products Continue to Be Contaminated With Deadly Fiber
The troubling news about the presence of asbestos in children's makeup is just the latest example of the deadly fiber contaminating imported products marketed toward children, said Environmental Working Group (EWG).
The national retail chain Claire's, which sells jewelry, makeup and other items targeted toward young girls and children, recently announced it was recalling a number of its makeup products after they tested positive for asbestos.
In 2015, EWG Action Fund (the 501 (c)(4) sister organization to EWG) found asbestos fibers in several brands of children's crayons and toy crime scene investigation kits.
In 2007, tests commissioned by the Asbestos Disease Awareness Organization (ADAO) also found the lethal fiber in another toy fingerprint named after the television show CSI: Crime Scene Investigation.
In 2000, an investigation by journalists with the Seattle Post Intelligencer discovered asbestos in imported crayons made with talc.
Shortly after EWG Action Fund published the results of its 2015 report, it called on the U.S. Consumer Product Safety Commission to investigate asbestos contamination in children's crayons and toys. EWG urged the CPSC to order toy companies to stop using talc that might be contaminated with asbestos in children's products.
"It seems every time someone looks for asbestos in imported children's products that include talc as an ingredient they find the deadly fiber," said Alex Formuzis, vice president for communications at EWG. "This should outrage both parents and federal regulators alike. There is no safe level of asbestos exposure so if a child inhales even the smallest amount it can lead to asbestos-triggered diseases, like mesothelioma, later in life."
All of the children's products where asbestos was detected were imported from China and included talc as an ingredient.
Geologically, talc and asbestos can be formed from the same parent rock. In many regions, talc deposits are contaminated with asbestos fibers.
Asbestos fibers lodged in the lungs or other organs can cause grave, often fatal, illnesses whose symptoms are not evident for decades after exposure. If children are exposed when young, there is more time for asbestos-related illness to develop later in life.
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Investors With Over $34 Trillion Demand Climate Crisis Action
Money talks. And today it had something to say about the impending global climate crisis.
Solar's Past, and Future
By Sam Cooper
Thomas Edison once said, "I'd put my money on the sun and solar energy. What a source of power!"
Zero Waste Kitchen Essentials
Simple swaps that cut down on kitchen trash.
By Kayla Robbins
Along with the bathroom, the kitchen is one of the most daunting areas to try and make zero waste.
Oil Leak Update: 1000x Worse Than Rig Owner Claimed, Still Going After 14 Years
The federal government is looking into the details from the longest running oil spill in U.S. history, and it's looking far worse than the oil rig owner let on, as The New York Times reported.
EcoWatch is pleased to announce our third photo contest!
'Up in Arms': New Book Explores the Bundys, Militias and the Battle Over Public Lands
By Tara Lohan
When armed militants with a grudge against the federal government seized the Malheur National Wildlife Refuge in rural Oregon back in the winter of 2016, I remember avoiding the news coverage. Part of me wanted to know what was happening, but each report I read — as the occupation stretched from days to weeks and the destruction grew — made me so angry it was hard to keep reading.
A searing heat wave has begun to spread across Europe, with Germany, France and Belgium experiencing extreme temperatures that are set to continue in the coming days.
In the 1980s, a Greenlandic subsistence hunter shot and killed a whale with bizarre features unlike any he had ever seen before. He knew something was unique about it, so he left its abnormally large skull on top of his toolshed where it rested until a visiting professor happened upon it a few years later.


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Benjamin Chavis Muhammad: An African American Civil Rights Leader Essay
Benjamin Chavis Muhammad: An African American Civil Rights Leader Essay
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Benjamin Chavis Muhammad is an African American civil rights leader. He was born on 22 January, 1948 in Oxford, North Carolina. He has taken his last name Muhammad sometime later in his life. His parents were Benjamin Chavis Sr. and Elisabeth Chavis. He was the only son of his parents among his three siblings. He did his schooling from the orphanage of colored people in North Carolina, where his mother worked as a teacher. Chavis became the member of National Association for Advancement of Colored People (NAACP) at the age of twelve. Chavis is married to Martha Rivera Chavis and has eight children.
Chavis served as an assistant to Dr. Martin Luther King at a very young age. As a result, he got inspired to work for Civil Rights Movement. At the age of thirteen, when he was still a wide-eyed boy, he made his first act of protest against racial prejudice. Moreover, Chavis was very influenced from his own family members, other than Dr. Martin Luther King. His great- great- grandfather, John Chavis, who was the first black person to graduate from Princeton University. John Chavis was a revolutionary solider; he was killed in 1838 for teaching slave children writing and reading.
At the age of 24, after being wrongly convicted of arson, Chavis gained international fame. He was sentenced for 34 years in prison and was freed in 1980. In 1988, he became the Vice President of National Council of Churches and in 1993, Chavis became the youngest CEO and executive director of NAACP. However, within sixteen months of joining NAACP, he was forced to resign after using some of NAACP fund to stop a sexual harassment lawsuit against him. In 1997, he joined Nation of Islam and adopted the surname Muhammad. He is also a cofounder of...
... middle of paper ...
...994). Chavis, Benjamin 1948–. Retrieved February 24, 2014, from Encyclopedia.com: http://www.encyclopedia.com/topic/Benjamin_Chavis_Muhammad.aspx
Jackson, E. (2011). Muhammad, Benjamin Chavis (1948- ). Retrieved February 24, 2014, from Black Past: http://www.blackpast.org/aah/muhammad-reverend-benjamin-chavis-1948
Jackson, G. (2005). Benjamin Chavis Muhammed. In We're Not Going to Take it Anymore (pp. 124-126). Beckham Publications Group, Inc.
Panel Members. (2014). Retrieved February 26, 2014, from The Brown & Black Presidential Forum: http://www.bbpresforum.org/panel.html
Psalms from Prison. (2014). Retrieved February 28, 2014, from AbeBooks: http://www.abebooks.com/9780829806618/Psalms-Prison-Chavis-Benjamin-Jr-082980661X/plp
The Brown & Black Presidential Forum. (2014). Panel Members. Retrieved February 26, 2014, from http://www.bbpresforum.org/panel.html
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Benjamin Franklin : The Greatest Thinkers, Inventors And Leaders Throughout American History
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