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Decay of FlhD and FlhC proteins in differentiating P. mirabilis harvested into LB alone or LB containing 5 mM dinitrophenol (DNP) or 5 mM sodium azide (NaN 3). Half-lives were determined as described in the legend to Fig. ​ Fig.2 2.Putative involvement of the Lon protease in FlhD and FlhC turnover. PCR1 and PCR2. a details the flhD operon and the two PCR reactions [ 11]. PCR1 is expected to yield a 1199bp product, PCR2 is expected to yield a 1344bp product in the absence of insertions or deletions. FP1stands for Forward Primer for PCR1, FP2 stands for Forward Primer for PCR2, RP stands for Reverse Primer for both PCR1 and PCR2. IS5 refers to the IS5 element at position −99 to −96 from the transcriptional start that was described by Barker et al. [ 11] and Wang and Wood [ 12]. IS1 refers to the IS1 elements that was described by Lee and Park [ 13] at −303 pb. IS refers to the IS5 described by Fahrner and Berg as the most distal (−476) IS element that had a positive effect on flhD expression [ 14]. b contains the 1kb ladder (Amresco, Solon OH) and 14 lanes of PCR products: lanes 1 & 2, MC1000; lanes 3 & 4, JS81; lanes 5 & 6, JS45; lanes 7 & 8, JS56; lanes 9 & 10, JS87; lanes 11 & 12, JS44; lanes 13 and 14, JS43. Odd numbered lanes show products of PCR1, even numbers lanes those of PCR2 Two model flagellar systems that form the foundation of the flagellar field are those from the enteric species E. coli and S. enterica. These two systems have led to key discoveries in relation to many aspects of flagellar structure, type 3 secretion, flagellar cell biology and the regulation of flagellar assembly. Textbook explanations suggest that most flagellar systems are being activated, regulated and built according to the models for E. coli and S. enterica. Modifications of transcriptional regulatory circuits contribute to the phenotypic diversity we see in closely related gene sets and we are only now able to investigate this in depth due to the tools available. Here we have taken a simple step and asked how do orthologous FlhD 4C 2 complexes function in the closely related species E. coli and S. enterica?

Samanta P, Clark ER, Knutson K, Horne SM, Prüß BM. OmpR and RcsB abolish temporal and spatial changes in expression of flhD in Escherichia coli biofilm. BMC Microbiol. 2013;13:182. The fine detail of the differences in the FlhD 4C 2 complexes only became apparent when we began to focus on their effect on flagellar gene expression and flagellar assembly. Biochemical analysis of isolated complexes showed that FlhC EC had weaker DNA binding ability to the P flgAB promoter region from S. enterica, consistent with previous investigations into FlhD 4C 2 DNA binding activity 19. The isolation of FlhD 4C 2 complexes from our strains suggested that a key aspect of the phenotypes we observed, was the stability of the complexes formed. Rappleye, C. A. & Roth, J. R. A Tn10 derivative (T-POP) for isolation of insertions with conditional (tetracycline-dependent) phenotypes. J Bacteriol 179, 5827–5834 (1997). Category III consisted of 30 isolates that produced PCR products larger than those of MC1000 in both reactions. Of the 30 isolates in this category, 28 isolates produced PCR fragments of 2 and 3.3 kb in PCR1 and PCR2, an example of this group is JS44 (Fig. 2b). This combination of PCR products is indicative of the continued presence of the IS5 from MC1000, supplemented by an insertion of approximately 800 bp within the part of the flhD operon that is amplified with the forward 1 primer. The remaining two category III isolates produced PCR1 and PCR2 products that were even larger. These are JS43 (Fig. 2b) and JS70, whose PCR products were 2.6 and 4 kb, respectively.Zhang Z, Saier Jr MH. Transposon-mediated adaptive and directed mutations and their potential evolutionary benefits. J Mol Microbiol Biotechnol. 2011;21:59–70. Chevance, F. F. V. & Hughes, K. T. Coordinating assembly of a bacterial macromolecular machine. Nat Rev Micro 6, 455–465 (2008). Field, Greg; Gantris, Peter; Gingerelli, Dain; Mitchel, Doug (2011). Harley-Davidson Buyer's Guide: 1984-2011. Minneapolis, MN US: MBI Publishing. ISBN 978-0-7603-3859-9 . Retrieved 2013-02-18. An 80cuin (1,300cc) engine was made optional on the Electra Glide in 1978, however, the FL designation was not changed. [14] By 1981, the 80cubicinch engine was the standard offering; the 74cuin (1,210cc) engine being discontinued. [16] Harley-Davidson Softail Springer Classic specifications". Archived from the original on 2007-07-05 . Retrieved 2008-08-22.

We allowed the highly motile E. coli K-12 strain MC1000 to form seven- and fourteen-day old biofilm, from which we recovered reduced motility isolates at a substantially greater frequency (5.4%) than from a similar experiment with planktonic bacteria (0.1%). Biofilms formed exclusively by MC1000 degraded after 2weeks. In contrast, biofilms initiated with a 1:1 ratio of MC1000 and its isogenic flhD:: kn mutant remained intact at 4weeks and the two strains remained in equilibrium for at least two weeks. These data imply that an ‘optimal’ biofilm may contain a mixture of motile and non-motile bacteria. Motility heterogeneity has been observed in other environments. One such environment is the mouse intestine. In the streptomycin-treated mouse intestine, flhD deletion mutants derived from the originally motile E. coli MG1655 parent eventually took over the population, but only after prolonged incubation [ 17, 18]. Intriguingly, 10 to 20% of the remaining bacteria had envZ missense mutations [ 19]. EnvZ is the histidine kinase of the osmoregulation system EnvZ/OmpR. The envZ P41L mutation increased the levels of phospho-OmpR [ 20], an inhibitor of flhD expression [ 21]. The authors concluded that the intestine likely contains niches where high flhD expression might be the advantage, alongside niches where low flhD expression might be an advantage [ 18]. The researchers did not attribute the entirety of this effect to motility. Instead, they proposed a ‘restaurant’ hypothesis [ 19], where the many metabolic genes that are regulated by flhD [ 22, 23] contribute to the niche adaptation. Our own studies with the aerotaxis sensor Aer, which impacts the sugar acid degradation pathway, support this hypothesis [ 24]. In 1966, the " Panhead" gave way to the " Shovelhead," gaining a ten percent increase in power in the process. [10] We inoculated 6 well polystyrene plates with MC1000, BP19, and mixtures of MC1000 and BP19 at ratios of 1:1, 10 MC1000/1 BP19 (designated 10:1), and 1 MC1000/10 BP19 (designated 1:10) in TB. Plates were covered with adhesive film (VWR International, Chicago IL) and incubated at 34°C. Biofilm amounts were determined after 1, 2, and 3weeks with the crystal violet assay (CV) [ 36]. Briefly, the planktonic bacteria were removed and the biofilms were washed twice with PBS, dried for 10min, and stained for 10min with 0.1% crystal violet in ddH 2O. The CV solution was removed and the biofilms were washed twice with PBS. CV was extracted from the bacteria with a mixture of 80% ethanol and 20% acetone, diluted 1:10, and the OD 600 was determined with a Synergy H1 plate reader from Biotek Instruments (Winooski, VT). Relative biofilm biomass is expressed as ratios where the OD 600 of each of the cultures was divided by that of MC1000. Averages and standard deviations were calculated across the three replicates for each strain/mixture. A Student’s t-test was used to determine the statistical significance of the difference between each mixture (or the non-motile mutant) and the MC1000 parent strain. A p-value below 0.05 was considered significant.Prüß BM, Campbell JW, Van Dyk TK, Zhu C, Kogan Y, Matsumura P. FlhD/FlhC is a regulator of anaerobic respiration and the Entner-Doudoroff pathway through induction of the methyl-accepting chemotaxis protein Aer. J Bacteriol. 2003;185:534–43. Other Softails with big forks followed, including the 1990 introduction of the FLSTF Fat Boy and the 1993-only FLSTN "Cow Glide". [28] [29] The base FLST Heritage Special was discontinued in 1991. [16] The FLSTN was continued after 1993 without the bovine motif as the Nostalgia model until 1996, it was replaced by the FLSTS Heritage Springer IN 1997. The Heritage Springer was produced through 2003, with the FLSTSC Springer Classic introduced in 2005. [16] The Heritage Springer and Springer Classic are designated as an FL despite using Springer forks instead of the large FL telescopic forks. [30] The first FL model, introduced in 1941, had the then-standard springer fork, with the teleskopic Hydraglide taking over from 1949 onwards. Tomoyasu, T. et al. The ClpXP ATP-dependent protease regulates flagellum synthesis in Salmonella enterica serovar typhimurium. J Bacteriol 184, 645–653 (2002). a b c d Field, Greg (2002). Original Harley-Davidson Panhead. St. Paul, MN US: MBI Publishing. p.27. ISBN 0-7603-1062-9.