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The ClpP protease homologue is required for the transmission traits and cell division of the pathogen Legionella pneumophila
© Li et al; licensee BioMed Central Ltd. 2010
Received: 29 September 2009
Accepted: 19 February 2010
Published: 19 February 2010
Legionella pneumophila, the intracellular bacterial pathogen that causes Legionnaires' disease, exhibit characteristic transmission traits such as elevated stress tolerance, shortened length and virulence during the transition from the replication phase to the transmission phase. ClpP, the catalytic core of the Clp proteolytic complex, is widely involved in many cellular processes via the regulation of intracellular protein quality.
In this study, we showed that ClpP was required for optimal growth of L. pneumophila at high temperatures and under several other stress conditions. We also observed that cells devoid of clpP exhibited cell elongation, incomplete cell division and compromised colony formation. Furthermore, we found that the clpP-deleted mutant was more resistant to sodium stress and failed to proliferate in the amoebae host Acanthamoeba castellanii.
The data present in this study illustrate that the ClpP protease homologue plays an important role in the expression of transmission traits and cell division of L. pneumophila, and further suggest a putative role of ClpP in virulence regulation.
Legionella pneumophila, a Gram-negative, intracellular bacterial pathogen, is the opportunistic agent responsible for a severe form of pneumonia named Legionnaires' disease and the less severe flu-like Pontiac fever [1, 2]. The remarkable capability of L. pneumophila to colonize a wide range of natural protozoa and mammalian host cells is mostly attributed to its unique Type IVB secretory system (T4BSS) whose components are encoded by the dot (defect in organelle trafficking) and icm (intracellular multiplication) genes [3–6]. L. pneumophila uses the Dot/Icm apparatus to inject effectors into the host cells to promote invasion and to modulate organelle trafficking, which in turn leads to formation of replication-permissive endosomes [7–9].
Similar to a variety of microbes, L. pneumophila undergoes a life cycle characterized by a biphasic conversion between a vegetative replicative form and a non-replicating, infectious and stress resistant transmissive form. On one hand, bacteria cultured in broth to either exponential or stationary phase display many similar attributes shared by the replicative and transmissive forms, respectively [10, 11]. For example, upon the transition from exponential phase to stationary phase, L. pneumophila becomes more infectious and more resistant to various stresses [12]. Furthermore, L. pneumophila in stationary phase also displays shortened cell body, flagellin expression, pigment accumulation and reduced sodium sensitivity. These attributes, together with virulence markers such as cytotoxicity, intracellular growth and phagocytosis, are recognized as the transmission traits of L. pneumophila[11, 13]. On the other hand, the in vitro-cultured stationary-phase L. pneumophila can achieve further differentiation to the cyst-like, hyper-infectious and resilient mature intracellular form (MIF) in aquatic environment or in specific mammalian cell lines. MIF is considered as an "in vivo stationary-phase form" while owning different outer membrane structure and protein composition compared with the stationary-phase form [14, 15]. In addition, an in vivo transcriptome of L. pneumophila was performed and exhibited the genes strongly induced in intracellular replicative or transmissive phase, respectively, which also revealed several virulence or transmission related genes specially induced intracellularly, confirming the dissimilarity between the in vitro- and in vivo- transmissive/stationary phase [16].
A complicated gene network has been implicated in the regulation of transmission traits in L. pneumophila. For example, the sigma factor RpoS, the two-component system LetA/LetS, and the quorum sensing regulator LqsR have all been shown to facilitate the expression of transmission traits [10, 11, 13, 17, 18]. CsrA, a global repressor of transmission [19], also appears to be tightly regulated by several factors such as PmrA (positive regulator of several Dot/Icm-translocated effector proteins) and rsm YZ (two non-coding RNAs) [20, 21]. In addition, CpxR has been found to activate transcription of several genes encoding components of the Dot/Icm complex as well as several Dot/Icm-translocated effectors [22, 23]. The concerted action of these regulators not only contributes to the display of transmission traits, but also plays a vital role in the re-entry into the replicative phase [11, 13, 19, 20, 24].
Proteolysis of detrimental and misfolded proteins is critically important for protein quality control and cellular homeostasis [25–27]. Four classes of energy-dependent protease systems have been identified throughout prokaryotes: ClpAP/XP, ClpYQ (also named HslUV), FtsH and Lon. ClpP and ClpQ, the catalytic cores of the proteases, require Clp ATPase chaperones for the recognition and unfolding of substrates; on the other hand, in FtsH and Lon, a single polypeptide contains both ATPase and proteolytic activity [26, 28]. The ClpP protease and Clp ATPase, which are widely distributed and highly conserved in various bacteria species as well as mitochondria and chloroplasts of eukaryotic cells [27, 29, 30], have been demonstrated to function in the regulation of stress response, sporulation and cell division [31, 32]. For example, ClpXP is responsible for the degradation of RpoS, the sigma regulator of stress response in E. coli[26]. In Salmonella enterica serovar typhimurium, loss of ClpXP has been shown to result in the over-expression of fliA and fliC, which in turn induced a hyperflagellate phenotype [33]. In Bacillus subtilis, ComK/S, the two-component regulator of competence and sporulation, are tightly controlled by the successive binding and degradation mediated by MecA and ClpCP [26]. ClpP also seems to regulate virulence in many pathogens such as Listeria monocytogenes, Streptococcus pneumoniae and Staphylococcus aureus[31, 34–36]. Finally, ClpP has been demonstrated to play a role in the biofilm formation [36–38].
As a ubiquitous bacterium in aquatic environment, L. pneumophila encounters numerous stresses such as elevated temperature, low pH and starvation during both planktonic existence and intracellular replication [11, 12]. We hypothesized that a rapid response to a changing environment might require an uncharacterized proteolytic system in L. pneumophila. In the present study, we explored the role of L. pneumophila ClpP in growth, stress tolerance, cell morphology and virulence to amoebae host. We demonstrate that ClpP affects several L. pneumophila transmission traits and cell division, and ClpP might play an important role in virulence regulation.
clpP homologue is required for optimal growth of L. pneumophila at high temperatures
clpP homologue is required for stress tolerance in stationary phase
clpP homologue is required for normal cell division of L. pneumophila
The combined results of SEM and cyro-TEM showed that unlike the "plump cocoid" shape of the WT or complemented strains, stationary-phase cells deficient in clpP were elongated and incapable to divide normally (Figure 4A to 4E). Furthermore, around 62% of LpΔclpP cells were twins, 23% were hyper-filamentous, and only 15% of cells were single (Figure 4F). In contrast, around 8% of WT JR32 cells were hyper-filamentous, and approximately 11% of cells were "twins" (Figure 4F). The abnormal cell morphology was also reversed by complementation (Figure 4C and 4F). These results together suggest that deletion of clpP lead to abnormal cell division and consequently aberrant cell morphology in L. pneumophila.
The LpΔclpP mutant is sodium tolerant
Loss of clpP impaires L. pneumophila growth and its cytotoxicity against A. castellanii
Cytotoxicity is an important virulent trait of L. pneumophila and correlates strongly with the function of the Dot/Icm T4SS [13, 44, 45, 47]. We next tested whether clpP homologue may affect the cytotoxicity of L. pneumophila against A. castellanii. L. pneumophila strains were used to infect A. castellanii with an MOI of 100. 24 h post infection, cytotoxicity was assayed by PI staining and quantified by flow cytometry analysis [13, 45]. As shown in Figure 6B, JR32 exhibited robust cytotoxicity (70% A. castellanii lethality), whereas LpΔclpP resulted in only 17% cell death, barely higher than that of the avirulent mutant ΔdotA (9% cell death). As expected, cytotoxicity was restored in the complemented strain LpΔclpP-pclpP (67% PI positive). These results were also confirmed by fluorescence microscopy (Figure 6C). Thus, it appeared that loss of clpP seriously impaires cytotoxicity against the amoebae host.
Loss of clpP abolishes intracellular multiplication of L. pneumophila in A. castellanii
Taken together, the adverse effects of clpP deletion on sodium tolerance, growth on APT, cytotoxicity and intracellular multiplication suggest that ClpP homologue might play an important role in virulence regulation of L. pneumophila.
In the current study, LpΔclpP was shown to exhibit reduced growth rate at high temperatures (Figure 2D) and impaired resistance to heat shock (Figure 3C) compared to the wild type. The LpΔclpP mutant also displayed impaired resistance to oxidative and low-pH conditions in stationary phase. As oxidative and acid stress are generally considered as harsh and detrimental to DNA [48, 49], ClpP homologue may play an important role in L. pneumophila DNA repair, consistent with its demonstrated function in E. coli[50], S. aureus[51] and Lactococcus lactis[52]. However, while several previous studies have demonstrated growth defect as a result of ClpP deficiency over a broad temperature range [34, 35, 51], deletion of clpP appeared to compromise the growth of L. pneumophila only at higher temperatures (Figure 2A to 2C), suggestive of a more restricted role independent of cold response.
Attenuation of ClpP or Clp ATPase activities has been shown to lead to abnormal bacterial morphology such as filamentation, aberrant cell wall structure and irregular cell division [29, 32, 53–55]. Likewise, results from SEM and cyro-TEM revealed that the LpΔclpP mutant cells were elongated and defective in cell division (Figure 4). Furthermore, SEM results also implicated a role of clpP in stress tolerance in L. pneumophila. In contrast to the defective cell surface observed in SEM (Figure 4D and 4E), largely normal cell surface were found by cyro-TEM in LpΔclpP mutant cells grown under normal conditions (Figure 4A to 4C), suggesting that the chemical treatment during SEM sample preparation, not clpP deletion, may have resulted in the abnormal cell surface.
How ClpP affects cell division is not fully understood. In C. crescentus, degradation of the cell cycle repressor CtrA by the ClpXP complex has been shown to contribute to G1-S transition, and deletion of clpP blocked cell division [54]. In B. subtilis, cells overproducing MurAA, an enzyme in peptidoglycan biosynthesis and a substrate of the Clp protease, displayed a filamentous, undivided morphology reminiscent of the clpP mutant cells, suggesting that degradation of MurAA by ClpP might contribute to normal cell segregation [56]. Furthermore, through a ClpP-independent pathway, the B. subtilis ClpX appeared to modulate the assembly of the tubulin-like protein FtsZ [57], which is known to be a key process in the replication and division of Gram-negative bacteria [58]. Identification of the substrate(s) for ClpP may shed light on the regulatory mechanism of cell division in L. pneumophila.
ClpP proteolytic complexes play pivotal roles in protein degradation or modification [26, 31, 32]. During the transition of B. subtilis cells to stationary phase, ClpP degrades massive amounts of proteins previously produced in exponential growth phase [32]. Notably, L. pneumophila also undergoes a biphasic life cycle with mutually exclusive gene expression for replication or transmission [10, 11]. While transiting from replication (exponential phase in vitro) to transmission (stationary phase in vitro), L. pneumophila activates an intricate network of regulators such as LetA/S, RpoS, PmrA, CpxR, rsm YZ, CsrA and LqsR [11, 13, 20, 21, 59]. As shown in our results, unlike the stationary-phase wild type which exhibits transmission traits, LpΔclpP mutant cells in stationary phase exhibit replicative forms such as reduced stress tolerance (Figure 2 and 3), cell elongation (Figure 4), enhanced sodium resistance (Figure 5), impaired cytotoxicity and growth on amoebae plates (Figure 6) and severely compromised intracellular multiplication in amoebae host (Figure 7). Thus, ClpP may play an important role in the transition from replication to transmission in L. pneumophila. On the other hand, several transmission traits are not affected by clpP-deletion such as pigment accumulation and transcription from the flaA (legionella flagellin coding) gene (our unpublished data), suggesting that the impact of ClpP on the transition to transmissive form in L. pneumophila is somewhat limited. Considering that ClpP always executes the post-transcriptional feedback regulation, and moreover, degrades the same substrates by cooperating with other proteases [26, 31], one explanation to such a limitation is that the degradation of ClpP substrates could be compensated by other proteases in clpP-deletion mutant, thus ClpP cannot govern the transition just as the global regulators such as RpoS, CsrA or LetA/S in L. pneumophila.
ClpP plays prominent roles in virulence of various Gram-positive pathogens such as S. aureus, S. pneumoniae and L. monocytogenes[34–36, 60]. Furthermore, ClpP was reported to control the levels of key virulence factors of type III secretory systems (T3SS) in certain pathogens such as S. typhimurium and Yersinia pestis[61, 62]. Recently, it was reported that loss of ClpP attenuated the virulence of Helicobacter pylori, a pathogen owning type IV secretory system (T4SS) [63]. It is interesting that clpP-deletion severely compromised the L. pneumophila infection against amoebae host (Figure 6 and 7). In our results, the sodium resistance exhibited by LpΔclpP mutant (Figure 5), which is a phenotype shared by the mutants without functional Dot/Icm T4SS [48, 64], together with the comparable decline in intracellular multiplication observed in LpΔclpP and ΔdotA mutants (Figure 7), suggest a role of ClpP in T4SS-dependent virulence through degrading a repressor or activating an up-regulator of the substrate(s) of ClpP. One possibility is that the ClpP protease has a major impact on the expression or function of Dot/Icm T4SS in L. pneumophila. Another possibility is that ClpP might be required for the expression of some T4SS substrates. In this case, loss of ClpP would also severely attenuate the intracellular growth even if the T4SS is intact, just as the case of L. pneumophila Sigma S factor (RpoS) [59]. Thus, identification of the substrate(s) of ClpP, which is currently underway in our laboratory, would help to discern the underlying relationship between ClpP and T4SS-dependent virulence in L. pneumophila.
In summary, our study shows that the L. pneumophila ClpP homologue is required for cell division and several transmission traits including stress tolerance, cell shortening, sodium sensitivity, cytotoxicity, growth on amoebae plates and intracellular multiplication. The study further suggests that the ClpP homologue might be important for virulence regulation of L. pneumophila.
Cells and reagents
Bacterial strains, plasmids and oligonucleotides used in this study.
Strain, plasmid or primer | Phenotype, genotype or sequence | Reference or source |
E . coli strains | ||
DH5α | F- endA1 hsdRI7 (rk - mk +) supE44 thi-1 λ- recA1 gyrA96 (Nalr) relA1 Δ (lacZYA-argF)U169 deoR φ 80dlacZ Δ M15 | Lab collection |
DH5αλpir | DH5α transduced with λpir | [69] |
L. pneumophila strains | ||
JR32 | Virulent L. pneumophila serogroup 1, strain Philadelphia, salt-sensitive isolate of AM511 | [43] |
LpΔclpP | JR32 with clpP deletion | This study |
LpΔclpP-pclpP | LpΔclpP containing pclpP | This study |
JR32-pBC | JR32 containing pBC(gfp)Pmip | This study |
LpΔclpP-pBC | LpΔclpP containing pBC(gfp)Pmip | This study |
LpΔdotA | JR32 with dotA deletion | Lab collection |
Plasmids | ||
pRE112 | Mobilizable suicide vector for construction of gene knockouts in G- bacteria, oriT oriV sacB Cm | [69] |
pMD18-T | cloning vector, Ap | TaKaRa |
pBC(gfp)Pmip | ColE1 ori Cm Pmip gfpmut2 | [70] |
pREΔclpP | pRE112::clpP for clpP deletion | This study |
pclpP | pBC(gfp)Pmip containing clpP under the control of mip promoter | This study |
Primers | ||
PXC-F1 | AGAGAGCTCCTGCCAGTAGGTCCTATAAG | This study |
PXC-R1 | TATGACATACAAGTTGCTGGACATTCTAC | This study |
PXC-F2 | CAACTTGTATGTCATAGGAACGCTCACC | This study |
PXC-R2 | GATGGTACCTGGGAAAATTGACAAACCGT | This study |
PXH-clpPF | TGGTGGAAGCTTTAGGAGTATCTAGCAAAGTTATAAGTC | This study |
PXH-clpPR | TGGTGGTCTAGATGAGAAAAAAGGAGAGTAAGC | This study |
DNA manipulation and chromosomal in-frame deletion
DNA manipulations were performed according to standard protocols [67]. All restriction enzymes were purchased from New England Biolabs. Pfu or Taq DNA polymerases were from TaKaRa. Purification of plasmids and genomic DNA was performed according to the manufacturer's instructions (Qiagen).
The in-frame deletion of clpP was performed by a non-polar strategy as described [68]. Briefly, upstream and downstream flanking sequences of clpP were amplified by PCR using the PXC-F1/PXC-R1 and PXC-F2/PXC-R2 primer pairs, respectively. The PCR products were mixed and then used as templates for the subsequent fusion PCR using the PXC-F1/PXC-R2 primers. Fusion PCR products were digested with Kpn I and Sac I and sub-cloned into the pRE112 suicide vector [69], yielding plasmid pREΔclpP. Allelic exchange was performed as follows. Briefly, pREΔclpP was introduced into the wild-type (WT) JR32 strain by electroporation and chloramphenicolR+ colonies were selected on BCYE-Cm plates. Transformants were inoculated into AYE and then incubated on BCYE containing 5% sucrose for 3 days at 37°C to select for strains devoid of the vector backbone. Positive colonies were confirmed by PCR and sequencing.
Complementation assay
A ColE1-type plasmid pBC(gfp)Pmip, carrying an enhanced GFP gene (gfpmut2) whose transcription was controlled by Pmip, the promoter of the Legionella-specific mip (macrophage infectivity potentiator) gene, was used for the clpP compensation experiment [70, 71]. As a control, the transcriptional activity of the mip promoter was not discernibly affected by the loss of clpP in JR32 (data not shown). pBC(gfp)Pmip was digested with Xba I and Hin dIII to remove the gfp. Sequences of clpP were amplified by PCR using the PXH-clpPF and PXH-clpPR primers, and the products were digested with Xba I and Hin dIII. The digestion products were ligated with the vector. The constructed plasmid pclpP was then electroporated into LpΔclpP, providing exogenous expression to compensate for the loss of clpP.
Growth experiments
The growth experiments were conducted using three L. pneumophila strains, including JR32 and the clpP deficient LpΔclpP derivative, both harboring the pBC(gfp)Pmip vector, as well as the complemented strain LpΔclpP-pclpP. These strains were first grown in 5 ml AYE for about 20 h. The cultures were expanded into 30 ml AYE in flasks, incubated to mid-exponential phase [optical density at 600 nm (OD600) 1.5-2.5], then diluted into new flasks to similar optical densities at approximate OD600 0.2. These new cultures were then incubated at 25°C, 30°C, 37°C, and 42°C, respectively. OD600 was determined by Beckman Du-530 at various time points.
Stress resistance assays
Resistance to stresses was measured as previously described [12, 65], with minor modifications. Cells from 1 ml broth cultures were centrifuged at 5,000 g for 5 min, and resuspended in AYE supplemented with 1 mM hydrogen peroxide, 0.1 M citric acid at pH 4.0, or 0.3 M potassium chloride, respectively. 30 min later (1 h for osmotolerance assay), cells were washed by centrifugation and resuspended in AYE. Cultures were subsequently serially diluted in water, plated on BCYE for colony forming unit (CFU) counting.
In heat resistance assays, cells from 1 ml broth cultures were centrifuged at 5, 000 g for 5 min and then resuspended in AYE. Samples for heat-shock were placed in a 57°C water bath for 20 min, with the control in a 37°C water bath. Cells were washed and serially diluted in AYE, and spread on BCYE for CFU counting.
Stress resistance was calculated as [(stressed sample CFU ml-1)/(control sample CFU ml-1)] × 100.
Sodium sensitivity assay
Sodium sensitivity assay was performed as previously described [65]. Briefly, cells from 1 ml broth cultures were centrifuged at 5, 000 g for 5 min and then resuspended in AYE. Subsequently, the cell suspensions were serially diluted in water, and spotted on BCYE and BCYE containing 100 mM NaCl or spread on plates for CFU counts. Sodium sensitivity was calculated as [(BCYE-100 mM NaCl CFU ml-1)/(BCYE CFU ml-1)] × 100.
Electron microscopy
For scanning electron microscopy (SEM), L. pneumophila cells in exponential or stationary phase were collected by centrifugation at 5,000 g for 2 minutes, and then washed 3 times with 1×PBS. After being fixed by 2% glutaraldehyde (pH 7.4) and 1% osmium tetroxide followed by dehydration in a graded ethanol series and isoamyl acetate embedding, the cells were dried by using a critical point drying method, and mounted on aluminum stubs and shadowed with gold. For visualization, a scanning electron microscope (Hitachi/Oxford S-520/INCA 300) was used at 10 kV.
For Cryo-transmisson electron microscopy, L. pneumophila cells were collected and washed using the same method as above. The cells were then resuspended in 1×PBS and 4 μl sample aliquots were directly applied to a holey carbon film grid (R3.5/1 Quantifoil Micro Tools GmbH, Jena, Germany), followed by blotting with filter paper (Whatman #1) for about 3 seconds. The grid was then immediately flash frozen by plunging into pre-cooled liquid ethane. The cryo-grid was held in a Gatan 626 Cryo-Holder (Gatan, USA) and transferred into TEM (JEOL JEM-2010 with 200 kv LaB6 filament) at -172°C. The sample was scanned and observed under minimal dose condition at -172°C. The micrographs were recorded by a Gatan 832 CCD camera at a nominal magnification of 10,000~ 50,000× and at the defocus of 3-5.46 μm.
Amoebae plate test (APT)
APT was performed as previously described [45]. Briefly, A. castellanii cells were cultured in PYG medium for 3 days prior to the test. A medium change was carried out one day before the test. The amoebae cells were washed off from the tissue culture flask, collected by centrifugation at 2,000 rpm for 5 min and resuspended in PYG to a density of 2 × 106 ml-1. 2 × 106A. castellanii cells were spread on BCYE agar plates, and incubated at room temperature overnight. Series of tenfold dilutions of stationary-phase bacterial cultures at a starting density of 1 OD600 (=109 cells ml-1) were prepared. 10 μl of each dilution were spotted onto the amoebae-CYET agar plates, and incubated at 37°C for 5 days.
Cytotoxicity assay using A. castellanii
To determine cytotoxicity, 2.5 × 105 amoebae cells were infected by bacteria at a multiplicity of infection (MOI) of 100. 24 h post infection, propidium iodide (PI) was added to 3 mg ml-1. A. castellanii cells were detached from the wells and 2.5 × 104 infected amoebae per sample were analyzed using a FACSCalibur flow cytometer (Becton Dickinson) with a scatter gate adjusted for A. castellanii[13]. Excitation was at 458 nm and fluorescence was measured at 495 nm. The data were collected and analyzed using the CELLQUEST software (Becton Dickinson). For fluorescence microscopy, the infected amoebae cells in each well of 24-well plates were stained with PI, then observed in bright field or by epifluorescence with an inverse microscope (Zeiss Axiovert 200 M, 20 × objective).
Intracellular growth in A. castellanii
For intracellular growth assays, exponentially growing A. castellanii were washed with Ac (A. castellanii) buffer, resuspended in HL5 medium, seeded onto a 24-well plate (2.5 × 105 per well) and were allowed to adhere for 1-2 h. L. pneumophila was grown for 21 h in AYE broth, diluted in HL5 and used to infect amoebae at an MOI of 10. The infection was synchronized by centrifugation at 440 g for 10 min, and the infected amoebae were incubated at 30°C. Thirty minutes post infection, extracellular bacteria were removed by washing 3 times with warm HL5 medium [13]. At the time points indicated, culture supernatant was removed and the amoebae cells were lysed with 0.04% Triton. The supernatant and the lysates were combined, and serial dilutions were prepared and aliquots were plated on CYE plates for CFU counting [72].
Statistical analysis
Basic statistical analyses were performed using Excel, and one-way ANOVA was performed using SPSS followed by a post hoc Student-Newman-Keul's test. The alignment of amino acid sequences was performed using the online ClustalW2 http://www.ebi.ac.uk/Tools/clustalw2.
- dot :
defect in organelle trafficking
- icm :
intracellular multiplication
- mip :
macrophage infectivity potentiator
mature intracellular form
multiplicity of infection
amoebae plate test
We thank Miss Ling-yan Zhu for kindly helping perform the flow cytometry analysis. This work was supported by the National Natural Science Foundation of China (No. 30670106, No. 30970123) and the Guangdong Provincial Natural Science Foundation of China (No.06201654) to YJL.
Authors' Affiliations
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- Gerth U, Krüger E, Derré I, Msadek T, Hecker M: Stress induction of the Bacillus subtilis clpP gene encoding a homologue of the proteolytic component of the Clp protease and the involvement of ClpP and ClpX in stress tolerance. Mol Microbiol. 1998, 28: 787-802. 10.1046/j.1365-2958.1998.00840.x.View ArticlePubMedGoogle Scholar
- Porankiewicz J, Wang J, Clarke AK: New insights into the ATP-dependent Clp protease: Escherichia coli and beyond. Mol Microbiol. 1999, 32: 449-458. 10.1046/j.1365-2958.1999.01357.x.View ArticlePubMedGoogle Scholar
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- Frees D, Savijoki K, Varmanen P, Ingmer H: Clp ATPases and ClpP proteolytic complexes regulate vital biological processes in low GC, Gram-positive bacteria. Mol Microbiol. 2007, 63: 1285-1295. 10.1111/j.1365-2958.2007.05598.x.View ArticlePubMedGoogle Scholar
- Tomoyasu T, Ohkishi T, Ukyo Y, Tokumitsu A, Takaya A, Suzuki M, Sekiya K, Matsui H, Kutsukake K, Yamamoto T: The ClpXP ATP-dependent protease regulates flagellum synthesis in Salmonella enterica serovar typhimurium. J Bacteriol. 2002, 184: 645-653. 10.1128/JB.184.3.645-653.2002.PubMed CentralView ArticlePubMedGoogle Scholar
- Gaillot O, Pellegrini E, Bregenholt S, Nair S, Berche P: The ClpP serine protease is essential for the intracellular parasitism and virulence of Listeria monocytogenes. Mol Microbiol. 2000, 35: 1286-1294. 10.1046/j.1365-2958.2000.01773.x.View ArticlePubMedGoogle Scholar
- Frees D, Qazi SN, Hill PJ, Ingmer H: Alternative roles of ClpX and ClpP in Staphylococcus aureus stress tolerance and virulence. Mol Microbiol. 2003, 48: 1565-1578. 10.1046/j.1365-2958.2003.03524.x.View ArticlePubMedGoogle Scholar
- Frees D, Chastanet A, Qazi S, Sorensen K, Hill P, Msadek T, Ingmer H: Clp ATPases are required for stress tolerance, intracellular replication and biofilm formation in Staphylococcus aureus . Mol Microbiol. 2004, 54: 1445-1462. 10.1111/j.1365-2958.2004.04368.x.View ArticlePubMedGoogle Scholar
- Lemos JA, Burne RA: Regulation and physiological significance of ClpC and ClpP in Streptococcus mutans. J Bacteriol. 2002, 184: 6357-6366. 10.1128/JB.184.22.6357-6366.2002.PubMed CentralView ArticlePubMedGoogle Scholar
- Wang C, Li M, Dong D, Wang J, Ren J, Otto M, Gao Q: Role of ClpP in biofilm formation and virulence of Staphylococcus epidermidis . Microbes Infect. 2007, 9: 1376-1383. 10.1016/j.micinf.2007.06.012.View ArticlePubMedGoogle Scholar
- Maurizi MR, Clark WP, Katayama Y, Rudikoff S, Pumphrey J, Bowers B, Gottesman S: Sequence and structure of ClpP, the proteolytic component of the ATP-dependent Clp protease of Escherichia coli . J Biol Chem. 1990, 265: 12536-12545.PubMedGoogle Scholar
- Wang J, Hartling JA, Flanagan JM: The structure of ClpP at 2.3 A resolution suggests a model for ATP-dependent proteolysis. Cell. 1997, 91: 447-456. 10.1016/S0092-8674(00)80431-6.View ArticlePubMedGoogle Scholar
- LeBlanc JJ, Davidson RJ, Hoffman PS: Compensatory functions of two alkyl hydroperoxide reductases in the oxidative defense system of Legionella pneumophila . J Bacteriol. 2006, 188: 6235-6244. 10.1128/JB.00635-06.PubMed CentralView ArticlePubMedGoogle Scholar
- Catrenich CE, Johnson W: Characterization of the selective inhibition of growth of virulent Legionella pneumophila by supplemented Mueller-Hinton medium. Infect Immun. 1989, 57: 1862-1864.PubMed CentralPubMedGoogle Scholar
- Sadosky AB, Wiater LA, Shuman HA: Identification of Legionella pneumophila genes required for growth within and killing of human macrophages. Infect Immun. 1993, 61: 5361-5373.PubMed CentralPubMedGoogle Scholar
- Byrne B, Swanson MS: Expression of Legionella pneumophila virulence traits in response to growth conditions. Infect Immun. 1998, 66: 3029-3034.PubMed CentralPubMedGoogle Scholar
- Albers U, Reus K, Shuman HA, Hilbi H: The amoebae plate test implicates a paralogue of lpxB in the interaction of Legionella pneumophila with Acanthamoeba castellanii. Microbiology. 2005, 151: 167-182. 10.1099/mic.0.27563-0.View ArticlePubMedGoogle Scholar
- Berger KH, Isberg RR: Two distinct defects in intracellular growth complemented by a single genetic locus in Legionella pneumophila . Mol Microbiol. 1993, 7: 7-19. 10.1111/j.1365-2958.1993.tb01092.x.View ArticlePubMedGoogle Scholar
- Vogel JP, Isberg RR: Cell biology of Legionella pneumophila . Curr Opin Microbiol. 1999, 2: 30-34. 10.1016/S1369-5274(99)80005-8.View ArticlePubMedGoogle Scholar
- Cooke MS, Evans MD, Dizdaroglu M, Lunec J: Oxidative DNA damage: mechanisms, mutation, and disease. FASEB J. 2003, 17: 1195-1214. 10.1096/fj.02-0752rev.View ArticlePubMedGoogle Scholar
- Xiao H, Li TK, Yang JM, Liu LF: Acidic pH induces topoisomerase II-mediated DNA damage. Natl Acad Sci USA. 2003, 100: 5205-5210. 10.1073/pnas.0935978100.View ArticleGoogle Scholar
- Gonzalez M, Rasulova F, Maurizi MR, Woodgate R: Subunit-specific degradation of the UmuD/D' heterodimer by the ClpXP protease: the role of trans recognition in UmuD' stability. EMBO J. 2000, 19: 5251-5258. 10.1093/emboj/19.19.5251.PubMed CentralView ArticlePubMedGoogle Scholar
- Michel A, Agerer F, Hauck CR, Herrmann M, Ullrich J, Hacker J, Ohlsen K: Global regulatory impact of ClpP protease of Staphylococcus aureus on regulons involved in virulence, oxidative stress response, autolysis, and DNA repair. J Bacteriol. 2006, 188: 5783-5796. 10.1128/JB.00074-06.PubMed CentralView ArticlePubMedGoogle Scholar
- Savijoki K, Ingmer H, Frees D, Vogensen FK, Palva A, Varmanen P: Heat and DNA damage induction of the LexA-like regulator HdiR from Lactococcus lactis is mediated by RecA and ClpP. Mol Microbiol. 2003, 50: 609-621. 10.1046/j.1365-2958.2003.03713.x.View ArticlePubMedGoogle Scholar
- Msadek T, Dartois V, Kunst F, Herbaud ML, Denizot F, Rapoport G: ClpP of Bacillus subtilis is required for competence development, motility, degradative enzyme synthesis, growth at high temperature and sporulation. Mol Microbiol. 1998, 27: 899-914. 10.1046/j.1365-2958.1998.00735.x.View ArticlePubMedGoogle Scholar
- Jenal U, Fuchs T: An essential protease involved in bacterial cell-cycle control. EMBO J. 1998, 17: 5658-5669. 10.1093/emboj/17.19.5658.PubMed CentralView ArticlePubMedGoogle Scholar
- Nair S, Poyart C, Beretti JL, Veiga-Fernandes H, Berche P, Trieu-Cuot P: Role of the Streptococcus agalactiae ClpP serine protease in heat-induced stress defence and growth arrest. Microbiology. 2003, 149: 407-417. 10.1099/mic.0.25783-0.View ArticlePubMedGoogle Scholar
- Kock H, Gerth U, Hecker M: MurAA, catalysing the first committed step in peptidoglycan biosynthesis, is a target of Clp-dependent proteolysis in Bacillus subtilis . Mol Microbiol. 2004, 51: 1087-1102. 10.1046/j.1365-2958.2003.03875.x.View ArticlePubMedGoogle Scholar
- Weart RB, Nakano S, Lane BE, Zuber P, Levin PA: The ClpX chaperone modulates assembly of the tubulin-like protein FtsZ. Mol Microbiol. 2005, 57: 238-249. 10.1111/j.1365-2958.2005.04673.x.View ArticlePubMedGoogle Scholar
- Margolin W: FtsZ and the division of prokaryotic cells and organelles. Nat Rev Mol Cell Biol. 2005, 6: 862-871. 10.1038/nrm1745.View ArticlePubMedGoogle Scholar
- Hovel-Miner G, Pampou S, Faucher SP, Clarke M, Morozova I, Morozov P, Russo JJ, Shuman HA, Kalachikov S: SigmaS controls multiple pathways associated with intracellular multiplication of Legionella pneumophila . J Bacteriol. 2009, 191: 2461-2473. 10.1128/JB.01578-08.PubMed CentralView ArticlePubMedGoogle Scholar
- Ibrahim YM, Kerr AR, Silva NA, Mitchell TJ: Contribution of the ATP-dependent protease ClpCP to the autolysis and virulence of Streptococcus pneumoniae. Infect Immun. 2005, 73: 730-740. 10.1128/IAI.73.2.730-740.2005.PubMed CentralView ArticlePubMedGoogle Scholar
- Hengge-Aronis R: Signal transduction and regulatory mechanisms involved in control of the sigma(S) (RpoS) subunit of RNA polymerase. Microbiol Mol Biol Rev. 2002, 66: 373-395. 10.1128/MMBR.66.3.373-395.2002.PubMed CentralView ArticlePubMedGoogle Scholar
- Jackson MW, Silva-Herzog E, Plano GV: The ATP-dependent ClpXP and Lon proteases regulate expression of the Yersinia pestis type III secretion system via regulated proteolysis of YmoA, a small histone-like protein. Mol Microbiol. 2004, 54: 1364-1378. 10.1111/j.1365-2958.2004.04353.x.View ArticlePubMedGoogle Scholar
- Loughlin MF, Arandhara V, Okolie C, Aldsworth TG, Jenks PJ: Helicobacter pylori mutants defective in the ClpP ATP-dependant protease and the chaperone ClpA display reduced macrophage and murine survival. Microb Pathog. 2009, 46: 53-57. 10.1016/j.micpath.2008.10.004.View ArticlePubMedGoogle Scholar
- Sexton JA, Miller JL, Yoneda A, Kehl-Fie TE, Vogel JP: Legionella pneumophila DotU and IcmF are required for stability of the Dot/Icm complex. Infect Immun. 2004, 72: 5983-5992. 10.1128/IAI.72.10.5983-5992.2004.PubMed CentralView ArticlePubMedGoogle Scholar
- Hammer BK, Swanson MS: Co-ordination of Legionella pneumophila virulence with entry into stationary phase by ppGpp. Mol Microbiol. 1999, 33: 721-731. 10.1046/j.1365-2958.1999.01519.x.View ArticlePubMedGoogle Scholar
- Segal G, Shuman HA: Legionella pneumophila utilizes the same genes to multiply within Acanthamoeba castellanii and human macrophages. Infect Immun. 1999, 67: 2117-2124.PubMed CentralPubMedGoogle Scholar
- Sambrook J, Russell DW: Molecular Cloning: a Laboratory Manual. 2001, Cold Spring Harbor, Cold Spring Harbor PressGoogle Scholar
- Ho SN, Hunt HD, Horton RM, Pullen JK, Pease LR: Site-directed mutagenesis by overlap extension using the polymerase chain reaction. Gene. 1989, 77: 51-59. 10.1016/0378-1119(89)90358-2.View ArticlePubMedGoogle Scholar
- Yu HB, Zhang YL, Lau YL, Yao F, Vilches S, Merino S, Tomas JM, Howard SP, Leung KY: Identification and characterization of putative virulence genes and gene clusters in Aeromonas hydrophila PPD134/91. Appl Environ Microbiol. 2005, 71: 4469-4477. 10.1128/AEM.71.8.4469-4477.2005.PubMed CentralView ArticlePubMedGoogle Scholar
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This article is published under license to BioMed Central Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.


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For the trail blazers who like a little speed in their step, the new GEL-FujiPro will make the perfect trail running companion. The low-profile Solyte midsole and Propulsion Trusstic® make for a spring-loaded ride, and the sleek Seamless Construction and Heel Clutching System are equal parts lightweight and fast. But the GEL-FujiPro means serious business out on the trail, complete with an Anti-Gravel Tongue, Trail Specific Outsole, and Lace Garage ideal for surging through the great outdoors. The shoe of choice for switching between trails and city streets Launch yourself forward with maximum efficiency thanks to Propulsion Trusstic One of the Fuji family, with great water release and lightweight set-up Feel lighter on every stride with the ultra-lightweight midsole
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subset: commoncrawl


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In quantum teleportation the sender, normally called Alice, instantaneously transfers the quantum state of a particle to a receiver, called Bob. In most experiments so far Alice has teleported the quantum state of a photon - defined in terms of its polarization - to Bob. The photon itself is not transferred: rather Bob's photon acquires exactly the same polarization as Alice's. The uncertainty principle means that Alice cannot know the exact state of her photon, but another feature of quantum mechanics called "entanglement" means that this is not an obstacle to teleporting the state to Bob.
Quantum entanglement essentially allows two particles to behave as one, regardless of how far apart they are. Photons can be entangled so that if one is vertically polarized, for instance, then the other photon in the pair is always horizontally polarized.
In a standard teleportation experiment a laser is directed at a crystal with nonlinear optical properties. Occasionally the photon will be "down-converted" into two lower energy photons, and sometimes these photons will have their polarizations entangled. In a teleportation experiment the beam is reflected back through the crystal to sometimes produce a second pair of entangled photons. By convention the photons in the first pair are labelled 2 and 3 (for mode 2 and mode 3), and those in the second pair are 1 and 4. Photons 1 and 2 are directed to Alice, photon 3 is sent to Bob and photon 4 is used as a trigger. A variety of mirrors, beamsplitters and polarizers are used to direct the photons to four detectors labelled D1, D2, D3 and T (for trigger).
The aim of the experiment is to transfer, or teleport, the polarization of photon 1 to photon 3. This is normally done by making a joint measurement on photons 1 and 2 which changes the polarization of the latter in such a way that photon 3 - which is entangled with it - always acquires the same polarization as the first photon. In other words the quantum state of photon 1 - which was unknown to Alice - has been teleported to photon 3.
The experiment was set up such that detectors D1, D2 and T all register photons at the same time when teleportation takes place. However, under certain circumstances - the emission of two photons each into modes 1 and 4 - it is possible for the same three detectors to register events even though there is no photon in mode 3 to teleport to. These spurious events mean that the D3 detector must also register - and destroy - a photon to be sure that teleportation has taken place.
By using a filter to reduce the intensity of the photons that are going to be teleported the researchers were able to significantly reduce the number of spurious detection events. The Vienna team could be 97% certain that the state had been teleported to photon 3 without actually having to detect it. Such a high accuracy means that the teleported photons could be used in "quantum repeaters" for long distance communication. The team now hopes to combine these results with a technique known as "entanglement purification" to further develop quantum communication over long distances.


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I needed to build up my stocks of comics and manga for my sons -- they're reading a lot of novels, so the novel-for-a-manga scheme is chugging along, and I also want to have a stash of comics for Christmas stockings -- so I placed a couple of mailorders for various comical funnies about a week ago, and they both arrived on Saturday.
(Anyone stalking me will have noted that I tweeted about the difficulty of getting into comics-shop packaging on that day.)
And, since I'm somewhat trying to rebuild my flooded graphic-novel collection (I have no hopes of rebuying any of the floppies; that's just a total loss), I got a few things for myself along the way:
Matt Wagner's Grendel Omnibus Volume 1: Hunter Rose -- I'm not crazy about the idea of this -- Devil by the Deed is a fine nasty little story, but all of the modern stylish crime junk Wagner has been weaving around it for the last decade-plus has obscured what made it special. So collecting it with all of those later Hunter Rose stories looks like a big mistake to me. But my copies of all of this stuff was turned to wet pulp, so you know what they say about beggars and choosers.
Elektra: Assassin by Frank Miller and Bill Sienkiewicz -- One of the great completely insane comics of our times, fizzing with livewire energy and pulsing with Miller's obsessions, then still (just barely) kept in check. I couldn't be without a copy of it for long.
Lobster Johnson Volume 2: The Burning Hand by Mike Mignola and various others -- I've gotten a year or two behind on my reading in the Hellboy-verse, for reasons that aren't entirely clear to me. I might have to read a whole bunch of it at once, sometime soon.
And Fables: Werewolves of the Heartland -- for a moment, I was excited, because the spine just says "Bill Willingham," and I thought maybe it was so late because he actually drew something, for the first time in a long time, but it's just corporate-comics being corporate; there are a whole slew of artists credited. I'm also solidly behind on Fables, and I probably should stop buying them until I read some.


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Now available in audio, originally published in Women on the Case, an anthology of 26 original stories by the best women in crime writing, a short story featuring Linda Barnes' feisty, redheaded, ex-cop private investigator, Carlotta Carlyle.
Carlotta can't stay away when she's offered a beautiful piece of classic blues history - the guitar of blues and gospel legend Reverend Gary Davis - in return for finding and stopping the uncomfortable advances of a blues-singing rock star's stalker.
©2013 Linda Barnes (P)2013 Brilliance Audio, Inc.


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These first couple of images were taken by an amateur photographer, J.J. Clarke (from County Monaghan) between 1890 and 1910 while a medical student in Dublin.
This is a shot of two fabulous ladies walking down Grafton Street in Dublin. This gives you a really good idea of what the fashion of the day was. Very formal and covered up!
The following is a photo taken in Bray (the Brighton of Ireland) of what ladies had to go through to go for a swim!
This was called a bathing machine and was strictly enforced for women at the time! Ladies had to enter the cabin and a horse pulled the cart into the water where ladies would emerge (fully clothed) for their seaside dip, modesty fully ensured! This was common practice in 19th century Europe and America but it continued to be used in certain Irish seaside resorts until the 1950s!
The next photo shows a Cincinnati suffragette dressed as a policewoman in 1909. The notion that women could possibly be employed in law enforcement was so alien that they had to dress up and present photographs so the public could grasp the idea! Note that the photo is marked -
"woman policeman
(ideal - not actual)"
(It was not until 1959 that Irish women were permitted to join An Garda Siochanna, with Mary Browne from Galway becoming the first Ban Gharda.)
Moving on to the 20's!
This photo shows a Washington policeman measuring the distance between a woman's knee and bathing suit in 1922. There was a law stating that bathing suits could not be over 6 inches above the knee.
And this photo shows what happened if you ignored the law. This was taken in Chicago in 1922, showing the arrest of several women daring to wear bathing suits that did not adhere to the legal hem length.
I could go on and on....!!!
Now that I am no longer spending every evening making vintage circus themed wedding paraphernalia I have a ridiculous amount of free time. With that in mind you can expect further posts from me on random subjects!!
x Caroline x


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Lot 4115: Pocket watch: very fine big and multicolored 20 K gold verge watch repeater, important Parisian master, Etienne Lenoir ca.1740-1760 (NO LIVE FEE)
123rd Cortrie Auction: Fine Pocket- & Wristwatches (NO LIVE FEE)
by Cortrie Spezial Auktionen
November 5, 2016
Hamburg, Hamburg, Germany
Live AuctionSold
Looking for the realized and estimated price?
Description: Ca. Ø 47 mm, ca. 95 g, 20 K gold, very splendid case, 4 different gold nuances, relief scene'palace' on back, floral tendrils on both sides, high-grade 5-part lateral hinge, fire-gilt verge movement, high-quality repeater, a toc, verge movement full signed, no.2006, original enamel dial in perfect condition, beautiful golden hands, minimal rubbed near hinge, altogether in excellent condition, very fine quality, in working order, one of the most important Parisian watchmakers of the 18th century.
Notes: LIVE FEE IS COVERED BY CORTRIE AUCTIONS.


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New Holland 461 Mower Conditioner Mower Conditioner Mower Conditioner Attachment Parts Manual |
Pages: 94 Weight: 0.50 lb. SKU: FO-P-460NHMC+ - 6078 |
FREE Expedited Priority Shipping (no Tax outside NJ) Price $42.99 |
This Parts Manual is for the New Holland 461 Mower Conditioner Attachment Mower Conditioner Mower Conditioner .This Parts Manual is critical for anyone who plans on servicing and repairing New Holland equipment.The Parts Manual includes 94 pages and weighs 0.50 pounds and ships via ground from our nearest US based warehouse. This manual is a new reproduction of the manuals printed in the time the machine was produced. Shipping to US 48 states only. 7 % sales tax New Jersey residents, no sales tax elsewhere. |
Tractorpoint Parts Fast Delivery! Low Prices Alternators |


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Glittered Wire Table Top Tree
Glittered Wire Table Top Tree
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Finish your room and table off in style this Christmas with a glittered wire bauble table top tree, great for creating that holiday spirit throughout the home. Mix with other decorations from the Heritage range for the perfect personalised look.
Technical Specifications
Colour description | Multicolour |
Finish | Glittered |
Pack quantity | 1 |
Weight | 66g |
Location | Internal |
Help & Advice
How to guides
How to personalise a Christmas wreath
The holidays are coming, so it's time to trim the tree and deck the halls. Festive decorations are an essential ingredient in the Christmas cocktail, and there are endless opportunities to get creative and make your home's seasonal style unique. At B&Q, we think one of the best ways to achieve a truly special festive look is by making your own decorations.
Finding a spare five minutes at Christmas time can be a challenge, but personalizing a festive wreath is a great way to create a unique decoration for your home and it's quick, easy and simple to make. Hang a Christmas wreath on your front door to impress your visitors, or place it centre stage on your dining table with an LED candle in the middle as a statement centrepiece.
Christmas wreaths are a festive favourite and with our step-by-step guide you'll be able to create one that's personal to your Christmas style.
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Proteins with unstable 3-D structures help the microscopic animals withstand drying.
The very cold, the merely chilled, and the colorful
The Science of Love, Bad Pharma, Genes, Cells and Brains, and Nature Wars
As cholera first tore through the Europe in the mid-19th century, people tried anything to prevent the deadly disease. Then science stepped in.
Meet some of the people featured in the February 2013 issue of The Scientist.
With dogged persistence and an unwillingness to entertain defeat, Bruce Beutler discovered a receptor that powers the innate immune response to infections—and earned his share of a Nobel Prize.
Six myths about job and salary negotiations and how they may hinder your ability to bargain effectively.
Can a vexing sense of entitlement actually aid in the pursuit of knowledge?
Because of their high protein and fat content and their reproductive efficiency, insects hold great promise for thwarting an impending global food crisis.
Tracking the genetic diversity and evolution of rhinoviruses can lead to a better understanding of viral evolution, the common cold, and more dangerous infections.


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A journey into peril.Cyrsania, high priestess of the Temple of Paladine, is blind to the beauty of the world but not to the joy of her faith. Lately, though, her communion with the god of light has become distant. Now a mysterious stone has come into her possession. Its power compels her to undertake a merilous quest, accompanied by a strange band of companions.Together they endure the burning heat of Kyrnn's most violet summer as they struggle toward their destination: the Temple at Neraka.
If they reach it, their world will be transformed forever.This novel, set in the time of the Chaos War, brings to life the story of Crysania, a Revered Daughter of Paladine, and Dalamar, the dark elf.
©1998 TSR, Inc. (P)2012 Audible, Inc.


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Have you ever purchased from PartSelect.com before?
Thank you for helping make our site better.
Below is a list of all parts for the Amana Dryer ALE643RAW - Electric. To find the repair part you need, you can browse the list below or you can narrow your choices by searching for a part by name or description.
This dryer drum glide pad is an inch long. It is made of a felt material and is used for gas and electric clothes dryers.
This 1/4 horsepower gas or electric clothes dryer motor carries 120 volts, 60 Hz, and 5.5 amps. It also performs 1725 revolutions per minute. This motor is used for some gas and some electric clothes dryers.
This part includes the upper and lower air duct assembly and lint screen.
Part may differ in appearance, but functions same as original part.
Sold individually. This bulb is a 7 watt appliance incandescent light bulb.This appliance light bulb is two inches long. It is used for many clothes dryers.
This drum support roller bracket and shaft assembly is a little over two inches long. It is made of metal.
This front cover spring retaining clip is not quite an inch long. It is made entirely of metal and is used for some washing machines.
This cycling thermostat has a limit of 153 degrees Fahrenheit, meaning that your dryer will cut out when the internal temperature reaches 153 degrees. It kicks back in though, when the temperature drops by 15 degrees Fahrenheit. This thermostat is just about two inches long.
This metal retaining ring is well under an inch long. Use it for your Maytag brand clothes washer or your clothes dryer.
This arm is what attaches to the idler pulley wheel.
Sold Individually.
This thermostat heater is two inches long. It is beige/brown in color and is made of plastic.
This clamp is used to secure the drive motor.


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This section contains 10,919 words (approx. 37 pages at 300 words per page) |
Dictionary of Literary Biography on Jean de La Bruyere
Jean de La Bruyère is known as the author of a single masterpiece, Les Caractères de Théophraste, traduits du grec, avec Les Caractères ou les mours de ce siècle (The Characters of Theophrastus, Translated from the Greek, with the Characters or the Mores of this Century), published in ever expanding editions from 1688 to 1696, the year of the author's sudden death at the age of fifty. It is a moral-satiric work attacking the vices of contemporary French society and made up primarily of portraits and maxims but accompanied by an extraordinarily diverse set of other types of discourse treating a wide range of subject matters. The whole work was intended, as he writes in the Préface, to describe the mours (meaning mores, customs, and forms of behavior as well as morals) of its original readers. The mirror...
This section contains 10,919 words (approx. 37 pages at 300 words per page) |


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Special Bulletin
The Patient Protection and Affordable Care Act ("Health Care Reform Act") was signed into law on March 23, 2010. Over the next several months, we will highlight major changes effectuated by this mammoth and historic legislation.
In this Special Bulletin, we will begin by addressing provisions which require employers to provide nursing mothers both time and space to express breast milk. We also will dispel some of the misconceptions and scuttlebutt associated with it.
What Is True And What Is Not True?
True: The Health Care Reform Act does include provisions that require employers who are subject to the Fair Labor Standards Act ("FLSA") to provide mothers reasonable break time to express milk for their infants who are up to one year old. The Health Care Reform Act also requires employers to provide a discreet location to perform the function (not a bathroom) which is shielded from view and free from intrusion from coworkers and the public.
The new provisions are added to Section 7 of the FLSA. Section 7 contains the basic time-and-one-half overtime provision and a grab bag of other wage and hour regulations.
Not True: It is not true that the effective date of the breast pumping provisions is delayed until September 23, 2010; January 1, 2011; or any of the other future effective dates provided for or contemplated by other parts of the Health Care Reform Act. These provisions became effective immediately on March 23, 2010. They are now the law of the land.
Not True: It is not true that the breast pumping provisions set up employers for a Title VII sex discrimination charge. They create a new duty, and employers therefore have perfect license to afford certain of their female employees a disparate term of employment (i.e., a mandated, albeit unpaid, break).
Not True: It is not true that employers with fewer than 50 employees are totally exempt from the breast pumping provisions. Instead, the Health Care Reform Act provides a limited exception for employers with fewer than 50 employees if the provisions "would impose an undue hardship by causing the employer significant difficulty or expense." Factors to be considered in making the "significant difficulty" determination are "the size, financial resources, nature or structure of the employer's business." The limited exception, according to its terms, applies to the time requirement, or the space requirement, or both, but there are many permutations for how the exception may play out in any given case.
The bar for compliance is set relatively low. For example, a small business that historically has made efficient use of space for operations will not have to add on a room to comply with the new requirements. An example that comes to mind would be a restaurant, whether fast food, casual, or elegant dining. Besides the areas for preparing food and serving customers, it may have only toilets for customers, perhaps separate toilets for employees, and a pantry for storage of inventory. The space constraints and low profit margins make strict compliance with the requirements impossible.
By their terms, the new provisions do not require space behind a locked door, only space that "is shielded from view and free from intrusion from coworkers and the public." An employer, therefore, could comply by installing a curtain and a sign reading "Keep Out When Closed." In other words, the space, "shielded from view and free from intrusion," may require not much more than something like a department store dressing room. Sensibility and sensitivity on this point may be informed by the not uncommon experience of seeing mothers nurse their babies in public, shielded from view by no more than a strategically placed shawl or blanket. Certainly, employers may be challenged, but the wording of the exception seems to invite the exercise of benign imagination.
What Employees Are Covered?
Because the breast pumping provisions have been enacted as an addition to the FLSA, consider the bigger picture in which the provisions fit. The FLSA sets minimum wage and overtime requirements on employers and employees engaged in interstate commerce (e.g., employees who manufacture, service, handle, ship, or receive goods which move across state lines, as well as employees who regularly cross state boundaries or routinely use the telephone or the mail while performing work). Also, all of a business's employees are deemed to be involved in interstate commerce if the business has some workers who produce, handle, work on, or sell goods that have been moved in, or were produced for, interstate commerce and the business has gross revenues in excess of $500,000. However, some businesses are always subject to the FLSA even if their gross revenues are less than the statutory minimum (viz., hospitals, rest homes, mental institutions, schools, colleges, and universities). The coverage of the FLSA is so wide and deep that employers who think they are exempt would do well to double check with legal counsel, unless their business is cooking free range pigs for friends and family off a country road in a mid-state county and never uses the telephone, couriers such as UPS or FedEx, or the U.S. mail.
A Concession To Employers
A concession to employers is found in the provision that employers are not required to compensate employees during a breast pumping break. Otherwise, breaks of 20 minutes or less normally would be required to be taken while on the clock.
For the breast pumping time to be non-compensable, however, the same rules that apply to bona fide meal time likely will have to be observed. Specifically, employees will have to be afforded sufficient time to effectively perform the act, completely relieved from duty. Nor would the "predominant benefit" test likely work in the case of breaks for breast pumping. The new provisions specifically require freedom from intrusion by coworkers. Employees should not be required to answer any questions, monitor calls, or experience similar work-related interruptions during a breast pumping break.
Potential Problems
Problems may arise in coordination of breast pumping breaks with other break time allowed by employers. One possible solution to anticipated inefficiencies would be a requirement that breast pumping breaks take the place of breaks otherwise allowed. However, it is important to recognize that while neither the FLSA nor the North Carolina Wage and Hour Act previously required employers to give adult employees any break time at all,1 employers now must allow for the breast pumping break.
Inefficiencies also can be expected during the months immediately post partum when a break for breast pumping may be required every couple of hours. The time needed for the break may vary as well, some methods being quicker than others. The new provisions provide for flexibility, using words and phrases such as "reasonable" and "each time such employee has need."
Additional Rules And Regulations
It is certain that the U.S. Department of Labor, which enforces the FLSA, will promulgate regulations and enforcement guidance. Phrases like "undue hardship" and its close companion "significant difficulty or expense" typically cause much regulatory and interpretive ink to be spilled.
The Health Care Reform Act also contains a provision that specifically allows the states to provide greater "protections" than those provided by the new breast pumping provisions. Neither North Carolina nor South Carolina has enacted any such law to date. There are, however, 24 states that currently have laws relating to breast pumping in the workplace, including Tennessee and Virginia. We can expect more to follow.
Employers on both sides of the 50-employee threshold will do well to review carefully issues of workplace time and space management with a magnanimous attitude of accommodating new mothers who return to work. Immediate attention should be given to employee handbooks that provide for employee break time, as these rules likely will have to be updated.
1North Carolina law does require breaks for youths under 16 years of age, who must be given at least a 30-minute break after five consecutive hours.
© 2010 Ward and Smith, P.A.
For further information regarding the issues described above, please contact William Joseph Austin, Jr.
This article is not intended to give, and should not be relied upon for, legal advice in any particular circumstance or fact situation. No action should be taken in reliance upon the information contained in this article without obtaining the advice of an attorney.
We are your established legal network with offices in Asheville, Greenville, New Bern, Raleigh, and Wilmington, NC.


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Wood, et al. v. Ionatron Inc.


text:
Test of Fire [Read by Dean Sluyter] Earth has been devastated by a massive solar flare. Now a small group of survivors fights to rebuild civilization. Cities became ovens, grasslands seas of flame. As the touch of dawn swept westward across the spinning planet, its fiery finger killed everything in its path. Glaciers in Switzerland began to melt; floodwaters poured down on burning Alpine villages. Paris became a torch, then London. North of the Arctic Circle, Laplanders in their summer furs burst into flame as their reindeer collapsed and roasted on the smoking tundra. The line of dawn raced westward across the Atlantic, but as it did, the sun dimmed as quickly as it had flared. The Americas escaped the sun's wrath . . . almost.
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text:
2013-03-25, 15:52 | Link #3 |
Senior Member
Join Date: Feb 2013
Location: Stockport UK
How many Anime series have a Character Album?
I was really impressed by the Sayonara Zetsubou Sensei one, especially because the songs were sung by the original cast & fitted each character well. I found some of the tracks on Youtube, & later bought the CD. |
2013-04-03, 19:25 | Link #7 |
Junior Member
Join Date: Aug 2012
Location: Midgard
Where did it say I expected there to be more Rock than Pop groups there?. I was wondering why there is such a small proportion of Rock in Anime, Didn't used to be that way. All the responses are wondering why I would ask, It seems I have joined the wrong forum. Mod can close topic now.
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