Surface smooth, rugose when old. Ostiolar dots Nutlin-3a chemical structure minute, olive or brown. Stroma colour first white, turning yellow, 4A3–4, brown-orange, 5CD5, greyish- to golden-yellow,

3B5–6, 4BC5–7, eventually (reddish) brown, 7E7–8, 6D7–8; mostly distinctly yellow when wet. Stromata when dry (0.6–)1.3–3.8(–8.0) × (0.4–)1.1–2.7(–4.7) mm, (0.3–)0.4–0.8(–1.1) mm thick (n = 75); solitary, gregarious or aggregated in small numbers (to 3) and pulvinate, or formed in large, subeffuse, flat and effluent, longish masses, becoming separated into individual stromata by cracks. Fertile part often flat, elevated on a short, stout, white stipe-like base, with margins laterally projecting beyond the base. Outline circular, angular, oblong or irregular. Margin margin sharply delimited, rounded, free, often white when young. Sides vertical

or slightly retracted downwards, find more white or yellowish, initially with radiating base mycelium. Surface initially typically with a white, later disintegrating, covering layer, smooth, finely granular to rugose, often slightly downy. Ostiolar dots minute, (20–)32–58(–80) μm (n = 75) diam, numerous, first often concealed by the covering white layer, becoming distinct, plane, less PF-02341066 in vivo commonly convex, with circular or oblong outline, brown. Stromata first of small white mycelial tufts, becoming compacted, turning argillaceous, pale to greyish yellow-orange, 4A3–4, 5A2–4, 4–6B4–5, 6A4, 6C4, mostly yellow with brown dots, i.e. yellow-brown, 5CD4–7, eventually pale brown to reddish brown, 6E6–8, 7CD5–6, 8E5–8, when old. Spore deposits white or yellow. Rehydrated mature stromata pulvinate, with plane, finely floccose, yellow surface and numerous distinct, plane, (orange-)brown ostiolar dots. Ostiolar openings hyaline in water. After addition of 3% KOH stroma

surface turning bright red to dark red; ostiolar openings hyaline; drying reddish brown. Immature stroma after rehydration semiglobose, smooth, white, with numerous irregular, plane or convex, almost light ochre dots; after addition of 3% KOH ostiolar dots first slightly orange, later surface turning homogeneously pale orange; eventually stroma macroscopically dark brown to nearly black. Stroma anatomy: Ostioles (58–)66–85(–92) μm long, plane or projecting to 20(–30) μm, (20–)25–40(–57) μm wide at the apex (n = 30), periphysate, sometimes with clavate marginal cells to 6 μm wide at the apex. Perithecia (148–)180–220(–230) × (90–)110–170(–205) μm (n = 30), 7–8 per mm stroma length, flask-shaped or globose, often crowded and laterally compressed; peridium (11–)14–17(–18) μm (n = 30) thick at the base, (3–)8–15(–18) μm (n = 30) thick at the sides, golden yellow; bright orange-red in 3% KOH.

g., maximum load, cortical volume, or cortical bone density. Fluid particle movement could also underlie the decreased fluoroscopy labeling at the endocortical surface observed in this study. Similar to Warden et al. [35], we hypothesize that a synergistic effect of the mechanotransduction

pathway in combination with muscle stimulation is responsible for the observations buy Fedratinib made here. Higher muscle activity results in increased bone formation, but these effects could be lower in comparison to WBVV at frequencies of 5–10 Hz. Garman et al. [38], who also observed an increase in trabecular bone after whole-body vibration, demonstrated that bone cells can detect physical stimuli directly in the absence of significant bone deformation. In their study, the oscillatory motion resulted in increased trabecular bone without altering weight bearing characteristics. A limitation of this study was the use of only one frequency, one direction of vibration, and one amplitude. this website The technique of WBVV used in this study was selected according to the results of Judex et al. [7], who demonstrated a significant increase of bone mass after WBV at 90 Hz compared to 45 Hz in rat tibiae. The results presented herein may not apply to subjects with older bones, nor may they apply to other bone regions, to males or even to humans. Our findings apply to a specific type of RSL3 supplier mechanical stimulus, and it is likely that other types

of vibration may result in varying effects on bone. Furthermore, rats were not fixed in a special position during vibration. In studies performed by Vershueren et al. [24] and Torvinen et al. [30], patients performed different actions during vibration. The test rats in this study moved freely on the vibration platform. It is possible that vibratory stimuli could change according to body posture. The effects could also potentially be dampened by the viscoelastic nature of the muscle–tendon apparatus [39]. In contrast to other groups that had animals laying

down on the vibration platform, the rats in this study tended to run all over the cage, attempting to escape from the cage by standing on their hind feet and thereby receiving greater axial load. The presented data and data from other studies suggest that mechanical signals may have the potential to influence both bone and muscle. Considering the mafosfamide importance of muscle strength and function to the incidence of falls and fall-related injuries, whole-body vertical vibration may be useful in reducing the risk for osteoporosis-related fractures [40]. Many questions remain regarding the benefit of whole-body vibration on the musculoskeletal system. It is not known, however, whether the effects will persist over time or whether such a treatment can help reduce falls and osteoporosis-associated fractures. Nevertheless, this non-drug method shows potential for the treatment of osteoporosis.

Many attempts have been made to find a definition of life that could be operational not only for terrestrial life, but for any form of “life” present in the universe, a definition Copanlisib solubility dmso that could help us to recognize a bona fide extraterrestrial “life” if we encounter it one day. In my opinion, such a definition is by essence biased by an idealist prejudice, reminiscent of Plato and Socrates’ ideas. It seems to imply that life is an ideal

form, concrete examples of life being various “shadows” of this ideal. I will adopt here the view that, up to now, life is only a terrestrial phenomenon, a characteristic of terrestrial “living organisms”. In fact, there is no life without living organisms and all presently known living organisms are thriving on planet Earth. If one day we hopefully meet friends from another world, it will then be possible to define “life” in term of the common properties shared by organisms from both planets. For the moment, the only materialistic way to define life is to start from the objects that exhibit

this extraordinary property: being alive (or having been EPZ5676 alive, once such objects are dead). In that sense, the question, “are BIBW2992 mw viruses alive?” is clearly at the heart of the debate. The answers to this question have varied in time, depending of our knowledge about viruses and our definition of life. Over the last decades, the answer has been often negative and viruses have been usually relegated to the periphery of the living world, being mainly considered as « dangerous » curiosities. They have been considered as by-products of cellular life, having probably originated as escaped genes from cellular organisms. However, this situation is rapidly changing, following

several discoveries made either by chance or by the effort of a few pioneers, and general advances in molecular biology (including the outcome of the genomic and post-genomic era) that have recently contributed to revise the position of viruses in the living world. Times are changing and viruses, once only considered as side-products of cellular Thymidine kinase evolution, are now at the center of many debates on the early evolution of life on our planet (Forterre 2002, 2005, 2006a, b; Brosius 2003; Bamford 2003; Bamford et al. 2006; Claverie 2006; Koonin et al. 2006; Ryan 2007; Raoult and Forterre 2008). Viral Particles Are the Most Abundant Biological Entities in the Biosphere It has been realized quite recently that viral particles are by far the most abundant biological entities on our planet (Suttle 2007). Indeed, they are ten times more abundant than bacterial cells in the upper ocean. This has been deduced in the nineties from examination of water samples by electron microscopy or epifluorescence optical microscopy.

It was suspected already in the mid-1970s and the early 1980s, and confirmed by later systematic LD spectroscopic studies, that the pigment dipoles are aligned under well-defined orientation angles with respect to the main axes of the complexes and/or of the membrane planes, and that the non-random orientation of the pigment molecules is a universal

property: this holds true for virtually all the photosynthetic pigments and in all organisms (Clayton 1980; Breton and Verméglio 1982). CD spectroscopy has also been widely used since the early 1970s and 1980s, during which the basic VRT752271 features and the occurrence of excitonic interactions in virtually all pigment–protein complexes have been established (Pearlstein 1991). In the last two decades, LD and CD spectroscopies

have gradually matured to become quantitative tools, which provide important information on different pigment systems and different, often high, levels of complexity, also under physiologically relevant conditions. Two chapters in the earlier books (Van Amerongen and Struve 1995; Garab 1996) have provided a detailed description of LD and CD techniques and the main areas of applications, while the monograph on photosynthetic excitons (Van Amerongen et al. 2000) has provided the theoretical background necessary for the in-depth interpretation of short-range, excitonic interactions between this website pigment molecules. For more complex, highly organized systems, the CD theory of psi (polymer and salt-induced)-type aggregates should be used (Keller and Bustamante 1986; Tinoco et al. 1987). The purpose of this educational review is to provide an introduction to LD and CD spectroscopies, as well as to some related differential

polarization spectroscopy and microscopy techniques. We explain, in simple terms, the basic physical principles and demonstrate, via Tyrosine-protein kinase BLK a few recent examples, the use of these tools in photosynthesis research. For a deeper understanding, readers are referred to the reviews and monographs cited above, and to articles quoted below in this review. For a basic understanding of the physical principles related to photosynthesis, see Clayton (1980). Transition dipole PRMT inhibitor moments of photosynthetic pigment molecules The absorption of light at a given wavelength corresponds to the transition from an electronic ground state to a given excited state of the molecule. The transition dipole moment, μ, which is associated with the electronic transition can be envisaged as a two-headed vector. The transitions for most photosynthetic pigments and most absorbance bands can be assigned to well-defined orientations with respect to the molecular coordinate system (see supplemental Fig. S1). (However, for some absorbance transitions, e.g.

The ompR transcription is induced directly by its own gene product in Salmonella enterica [3]. OmpR consensus-like sequences are found in the upstream region of ompR in Escherichia coli, although there are still no reported experimental data for its autoregulation in this bacterium. Upon the elevation of medium osmolarity, cellular OmpR-P levels are likely enhanced by two distinct mechanisms, namely, post-translational phosphorylation/dephosphorylation by EnvZ and transcriptional auto-stimulation. Enterobacteriaceae express at least two major outer membrane (OM) porins, namely,

OmpF and OmpC, both of which form transmembrane pore structures and function as ion channel [4–6]. OmpF and OmpC in the cell of E. coli form water-filled pores that are poorly selective selleck screening library to cations (so called non-specific porins), thereby allowing the diffusion of low-molecular-weight polar compounds (not over 600 daltons) into the cell to maintain cell permeability. They exist as homotrimers in the OM. The basic structural element of the porin monomer is an ellipsoid in the section cylinder consisting of 16 transmembrane β-strands (so-called β-barrel)

connected by short periplasmic and longer ‘external’ A-1210477 loops [7]. E. coli OmpX contains 8-stranded β-barrel, with polar residues on the inside and hydrophobic residues on the outside facing the membrane environment [8]. Enterobacter aerogenes OmpX is the smallest known channel protein with a markedly cationic selectivity [6, 9, 10]. Although several experiments have demonstrated that OmpX plays check details roles that are similar to those of porin [6, 9–12], it is not yet clear whether or not OmpX forms this website porins on the

cell membrane. E. aerogenes OmpX forms channels in the lipid bilayer [6]; however, the NMR and crystal structures of OmpX do not show pores [8, 13]. The ompX expression in E. coli [12] or E. aerogenes [6] is enhanced during early exposure to environmental perturbations, such as high osmolarity, antibiotics and toxic compounds, that are accompanied by the repressed expression of non-specific porins (OmpF and/or OmpC). Over-expression of OmpX, with a channel structure that is much smaller than that of OmpF and OmpC [6], may stabilize cell OM and balance the decreased expression of the two non-specific porins for the exclusion of small harmful molecules. It is interesting to further investigate the roles of OmpX in modulating OM permeability and adaptability. OmpR consensus-like sequences have been found within the ompX upstream region in E. coli and E. aerogenes [6]; however, the regulation of ompX by OmpR has not yet been established experimentally in any bacterium. As shown in E. coli as a model, OmpF and OmpC are reciprocally regulated by medium osmolarity.

Otherwise, ex situ activities for an increasing number of threatened species, other than a handful of charismatic mega vertebrates, are inevitably destined to fail. Acknowledgments I wished to thank several colleagues for sharing ideas and opinions; DMXAA ic50 they are

C. Lees, J.-M. Lernould, A. Kitchener, H. Schram, K. Kawata, and R. Wirth. References Amori G, Gippoliti S (2000) What do mammalogists want to save? Ten years of mammalian conservation biology. Biodivers Conserv 9:785–793CrossRef Anderegg R, Frey H, Muller HU (1984) Reintroduction of the bearded vulture or lammergeyer (Gypaetus barbatus aureus) to the Alps. Int Zoo Yearb 23:35–41 Backer A (2007) Animal ambassadors: an analysis of the effectiveness and conservation impact of ex situ breeding efforts. In: Zimmermann A, Hatchwell M, Dickie L, West C (eds) Zoos in the 21st century. Catalyst for conservation? Cambridge University Press, pp 139–154 Balmford A, Mace GM, Leader-Williams N (1996) Designing the ark: setting priorities for captive breeding. Conserv Biol 10:719–727CrossRef Barnett R, Yamaguchi N, Barnes I, Cooper

A (2006) Lost populations and preserving genetic diversity in the lion Panthera leo: Trichostatin A ic50 implications for its ex situ conservation. Conserv Genet. doi:10.​1007/​s10592-005-9062-0 Bowkett AE (2009) Recent captive-breeding EPZ004777 solubility dmso proposals and the return of the ark concept to global species conservation. Conserv Biol 23:773–776CrossRef Brito D, Oprea M (2009) Mismatch of research effort and conservation in avian conservation biology. Trop Conserv Sci 2:353–362 Burger J, Hemmer H (2006) Urgent call for further breeding of the relic zoo population of the critically endangered barbary lion (Panthera leo leo Linnaeus 1758). Eur J Wildl Res 52:54–58CrossRef Calvignac S, Hughes S, Hanni Amrubicin C (2009) Genetic diversity of endangered brown bear (Ursus arctos) populations at the crossroads of Europe, Asia and Africa. Diver Distrib 1–9. doi:10.​1111/​j.​1472-4642.​2009.​00586.​x Conde DA, Flessness

N, Colchero F, Jones OR, Scheuerlein A (2011) An emerging role of zoos to conserve biodiversity. Science 331:1390–1391PubMedCrossRef Conway W (2007) Entering the 21st century. In: Zimmermann A, Hatchwell M, Dickie L, West C (eds) Zoos in the 21st century. Catalyst for conservation? Cambridge University Press, Cambridge, pp 12–21 Conway W (2011) Buying time for wild animals with zoos. Zoo Biol 30:1–8PubMed Durrell L, Anderson DE, Katz AS, Gibson D, Welch CR, Sargent EL, Porton I (2007) The Madagascar fauna group: what zoo cooperation can do for conservation. In: Zimmermann A, Hatchwell M., Dickie L, West C (eds) Zoos in the 21st century. Catalyst for conservation? Cambridge University Press, Cambridge, pp 275–286 Frynta D, Lišková S, Bültmann S, Burda H (2010) Being attractive brings advantages: the case of parrot species in captivity. PLoS ONE 5(9):e12568. doi:10.​1371/​journal.​pone.

We identified these two spots as the HSP60, with a molecular weight of 61.055 kDa, and a pI value of 5.70. AZD7762 concentration The spectrum figure of HSP60 was presented in Figure 2. Western blot results using the cell Bioactive Compound Library cell assay lysates samples confirmed the findings that the expression of HSP60 was significantly lower in the cell lysates of PcDNA3.1(IGFBP7)-RKO transfectants. A representative image was presented in Figure 3A. The

secretion of HSP60 was also compared between the supernatants from PcDNA3.1(IGFBP7)-RKO transfectants and controls using ELISA. Secretion of HSP60 was also found to be downregulated by IGFBP7 (Figure 3B). Figure 2 LC-MS spectrum obtained for spot 9. Peptide fragments were analyzed by LC-tandem MS and MALDI-TOF analysis SN-38 of peak m/z was performed. Major monoisotopic peaks of trypsin-digested peptides, detected by MALDI-TOF MS, are indicated on the spectrum. The sequence of HSP60 protein was represented by single-letter code for amino acids on the top left corner of the image, where peptide matches between the sample and the HSP60 sequence are shown bold. Figure 3 Downregulation of HSP60 protein expression in PcDNA3.1( IGFBP7 )-RKO transfectants. A: Whole cell lysates of the stable PcDNA3.1(IGFBP7)-RKO transfectants and PcDNA3.1-RKO transfectants were prepared,

and equal amounts of protein (50 μg/lane) were loaded. HSP60 expression Methamphetamine was assessed using a rabbit anti-HSP60 antibody. GAPDH is used as an internal loading control. Shown is representive of experiments performed on at least three different isolations. The amount of HSP60 protein expression in PcDNA3.1(IGFBP7)-RKO transfectants was lower than that of the control group. B: HSP60 concentration in cell supernatants was measured using the HSP60 ELISA kit according to the manufactures instructions. Experiments were performed in triplicates. Results represent the mean HSP60 concentration (ng/ml) ± SD,. *, p < 0:05 vs. control. Recombinant HSP60 reversed the proliferation inhibition induced by IGFBP7 To

clarify the biological effect of HSP60 downregulation induced by IGFBP7 in RKO cells, we studied the function of recombinant HSP60 on the proliferation of PcDNA3.1(IGFBP7)-RKO cells. We found that addition of HSP60 protein could promote the cell proliferation rate of PcDNA3.1(IGFBP7)-RKO cells(Figure 4A). HSP60 could also increase the colony formation ability and the colony size of the cells (Figure 4B, 4C). Figure 4 HSP60 protein decreased the proliferation rate and colony formation ability of PcDNA3.1( IGFBP7 )-RKO cells. A: PcDNA3.1(IGFBP7)-RKO cells were plated in sextuple in 96-well microtitre plates at 3 × 103/well, cultured with medium with or without recombinant HSP60(1 μg/ml). Ten μl of CCK8 was added to each well at the indicated time (12 h, 24 h, 36 h, 48 h, 60 h, 72 h).

Figure 3 Capacitance-voltage curves (a) and current–voltage characteristics (b) of Al/Er 2 O 3 /TaN and Al/Er 2 TiO 5 /TaN structure devices.

The transfer characteristics of the a-IGZO TFT devices using Er2O3 and Er2TiO5 gate dielectrics were shown in Figure  4a. The V TH value of the Er2O3 and Er2TiO5 a-IGZO TFT devices is 1.5 and 0.39 V, whereas the I on/I off ratio is 1.72 × 106 and 4.23 × 107, respectively. The moisture absorption of the Er2O3 film generates a rough surface due to the formation of Er(OH) x , thus causing degradation in the electrical characteristics. Furthermore, the I off current can be improved by bottom gate pattern to reduce the leakage path from the gate to the source and drain. Furthermore, the μ FE of the Er2O3 and Er2TiO5 TFT devices is 6.7 and 8.8 cm2/Vs. This

result is due to the smooth roughness at the oxide-channel interface [15]. The subthreshold PARP inhibitor swing (SS) of the Er2O3 and Er2TiO5 TFT devices is 315 and 143 mV/dec, respectively. The titanium atoms can effectively passivate the oxygen vacancies in the Er2TiO5. The effective interface trap state densities (N it) near/at the interface between the STI571 nmr dielectric and IGZO were estimated from the SS values. By neglecting the depletion capacitance in the active layer, the N it can be calculated from the relationship [6]: (1) where q is the GSI-IX research buy electronic charge; k, the Boltzmann’s constant; T, the temperature; and C ox, the gate capacitance density. The N it values of IGZO TFTs using Er2O3 and Er2TiO5 gate dielectrics are about 6.92 × 1012 and 2.58 × 1012 cm−2, respectively. Figure  4b shows the output characteristics of the a-IGZO TFT devices using the Er2O3 and Er2TiO5 gate dielectrics. As is seen, the driving current increases significantly for the

Er2TiO5 dielectric material. This outcome may be attributed to the higher mobility and smaller threshold voltage. Figure 4 Transfer and output characteristics. Transfer characteristics (I DS-V GS) (a) and output characteristics (I DS-V DS) (b) of high-κ Er2O3 and Er2TiO5 a-IGZO TFT devices. Urease To explore the reliability of an a-IGZO transistor, the dc voltage was applied to the high-κ Er2O3 and Er2TiO5 a-IGZO TFT devices. Figure  5a shows the threshold voltage and drive current degradation as a function of stress time. The voltage stress was performed at V GS = 6 V and V DS = 6 V for 1,000 s. The shift in threshold voltage and the degradation in drive current are associated with the trap states in the dielectric layer and the interface between the dielectric film and channel layer [16]. The large V TH shift (1.47 V) of the Er2O3 TFT can be due to more electrons trapping near/at the interface between the Er2O3 and IGZO layer [6], whereas the low V TH shift (0.51 V) of the Er2TiO5 TFT device may be attributed to the reduction of the trapped charge in the film.

Similar results were observed with the in vivo experiments as well. Although fewer pups died within 24 hrs post-infection in the groups infected with RS218cured as compared to the groups infected with wtRS218 and RS218compl, there was no statistically significant difference in mortality rates between the three groups (Figure 5B). No mortalities were detected in the negative control group treated with

PBS or E. coli DH5α. In groups infected with wtRS218 or RS218compl, 84-87% of rat Vistusertib clinical trial pups that survived 24 hrs post-infection showed septicemia, whereas in groups treated with RS218cured strain, only 33% had septicemia. In all three groups the number of bacteria in the blood was too numerous to count (>1.5-2.8 *104 CFU/ml). Also, E. coli were re-isolated buy NVP-BSK805 from CSF collected from 84-87% of rat pups infected with wtRS218 or RS218compl whereas only 29% CSF samples collected from rat pups infected with RS218cured strain contained E. coli suggesting a role of pRS218 in translocation of bacteria through the blood brain barrier (BBB) to cause meningitis. Similarly, histopathological

evaluation of brain tissue from the rat pups inoculated with wRS218 or RS218compl strains demonstrated lesions consistent with meningitis (Figure 6). The bacterial loads in CSF were 4.57 + 3.02*103 in rat pups infected with wtRS218 strain and 3.77 + 2.24*103 in rat pups infected with RS218cured strain. Figure 4 Confirmation of pRS218 curing. A, Plasmid profiles of wtRS218 and RS218cured. B, PCR amplification of selected pRS218 genes in wtRS218 and RS218cured. Lane 1,100 bp Erismodegib order ladder; Lane 2, senB; Lane 3, scsD; Lane 4, transposase; Lane 5, traU; Lane 6, pRS218_113; Lane 7, ycfA; Lane 8, ompA. C, during Growth of wtRS218 and RS218cured E. coli in LB broth, M9 medium containing 10 μg/ml niacin broth (M9),

and complete cell culture medium (CM). Figure 5 Evaluation of virulence potential of pRS218 in vitro and in vivo. A, Involvement of pRS218 in invasion of hCMEC cells. B, Comparison of mortality, septicemia and meningitis among the groups of rat pups infected with wtRS218, RS218cured, RS218compl. ** denotes statistical significance and * denotes no statistical significance. Figure 6 Histopathological evaluation of brain tissue from rat pups. Five-day-old rat pups were infected by the IP route with wtRS218, RS218compl, RS218cured, E. coli DH5α or PBS. Pups that survived were euthanized 24 hrs post-infection, and the brains were excised, embedded in formalin, sectioned in paraffin, and stained with haematoxylin and eosin. A-F: meningitic lesions observed in pups infected with wtRS218 (A and B) or RS218compl (C, D, E, and F). Arrows indicate rod-shaped bacteria in meninges and brain tissue (black), neutrophilic infiltration/neutrophilia (blue), and cerebral edema (orange). G to I: normal histology of brain tissue from pups inoculated with RS218cured (G), PBS (H) or DH5α (I).

Members of the P. syringae species are gram negative plant-associated γ-proteobacteria that can exist both as harmless epiphytes and find more as pathogens of major agricultural crops [48–52]. Pathogenic varieties of this species utilize a Hrc-Hrp1 T3SS to inject effector proteins and thus subvert signalling pathways of their plant hosts. This secretion system (Hrc-Hrp1 T3SS) and its effector proteins are responsible for the development of the characteristic disease symptoms on susceptible plants and the triggering of the Hypersensitive Response (HR) in resistant plants [26, 49, 50, 52]. Comparative genomics of closely related

isolates or species of pathogenic bacteria provides a powerful tool for rapid identification of genes involved in host specificity and virulence [53]. In this work, we reported sequence Selleck Screening Library similarity searches, phylogeny analysis and prediction

of the physicochemical characteristics of the hypothetical T3SS-2 proteins, as well as gene synteny analysis of the BGB324 clinical trial T3SS-2 gene cluster in P. syringae pv phaseolicola 1448a, P. syringae pv oryzae str. 1_6 and P. syringae pv tabaci ATCC11528 in order to characterize this recently identified gene cluster. This analysis revealed that the T3SS-2 most closely resembles the T3SS of the Rhc-T3SS family. It further typifies a second discrete subfamily (subgroup II) within the Rhc-T3SS family in addition to the ones represented by the R. etli T3SS (subgroup III) and the known Rhizobium-T3SS (subgroup I). Usually, the presence of two T3SS gene clusters in the same genome is not the result of gene duplication inside the species

but rather the result of independent horizontal gene transfers. This may reflect progressive coevolution of the plant patho/symbio-system to either colonize various hosts or interact with the plant in different disease/symbiotic Rho stages. In our phylogenetic analysis proteins encoded in the T3SS-2 cluster of P. syringae strains are grouped together with the Rhizobium NGR234 T3SS-2. This finding suggests the possibility of an ancient acquisition from a common ancestor for Rhizobium NGR234 T3SS-2 and the P. syringae T3SS-2. T3SSs of the Rhizobium family possesses a GC-content in same range (59-62%), a value lower than the chromosome average. Since the GC content of T3SS-2 is almost the same as that of the genome of the P. syringae strains, it is difficult to characterize the second T3SS gene cluster as a genomic island based solely on this criterion. However, the genome sequencing of two other members of P. syringae [pathovars tomato DC3000, syringae B728A] revealed the total absence of a T3SS-2 like cluster. The T3SS-2 gene cluster found in P. syringae pv phaseolicola 1448a, P. syringae pv oryzae str.1_6, P. syringae pv tabaci and of Rhizobium sp. NGR234, is also present in P. syringae pv aesculi (strains NCPPB 3681 and 2250)[54], P. syringae pv savastanoi (str. NCPPB 3335) [55], P.