EPEC mutant strains generated with the marker-less deletion strategy.
\\n\\n
Dr. Pletser’s experience includes 30 years of working with the European Space Agency as a Senior Physicist/Engineer and coordinating their parabolic flight campaigns, and he is the Guinness World Record holder for the most number of aircraft flown (12) in parabolas, personally logging more than 7,300 parabolas.
\\n\\nSeeing the 5,000th book published makes us at the same time proud, happy, humble, and grateful. This is a great opportunity to stop and celebrate what we have done so far, but is also an opportunity to engage even more, grow, and succeed. It wouldn't be possible to get here without the synergy of team members’ hard work and authors and editors who devote time and their expertise into Open Access book publishing with us.
\\n\\nOver these years, we have gone from pioneering the scientific Open Access book publishing field to being the world’s largest Open Access book publisher. Nonetheless, our vision has remained the same: to meet the challenges of making relevant knowledge available to the worldwide community under the Open Access model.
\\n\\nWe are excited about the present, and we look forward to sharing many more successes in the future.
\\n\\nThank you all for being part of the journey. 5,000 times thank you!
\\n\\nNow with 5,000 titles available Open Access, which one will you read next?
\\n\\nRead, share and download for free: https://www.intechopen.com/books
\\n\\n\\n\\n
\\n"}]',published:!0,mainMedia:null},components:[{type:"htmlEditorComponent",content:'
Preparation of Space Experiments edited by international leading expert Dr. Vladimir Pletser, Director of Space Training Operations at Blue Abyss is the 5,000th Open Access book published by IntechOpen and our milestone publication!
\n\n"This book presents some of the current trends in space microgravity research. The eleven chapters introduce various facets of space research in physical sciences, human physiology and technology developed using the microgravity environment not only to improve our fundamental understanding in these domains but also to adapt this new knowledge for application on earth." says the editor. Listen what else Dr. Pletser has to say...
\n\n\n\nDr. Pletser’s experience includes 30 years of working with the European Space Agency as a Senior Physicist/Engineer and coordinating their parabolic flight campaigns, and he is the Guinness World Record holder for the most number of aircraft flown (12) in parabolas, personally logging more than 7,300 parabolas.
\n\nSeeing the 5,000th book published makes us at the same time proud, happy, humble, and grateful. This is a great opportunity to stop and celebrate what we have done so far, but is also an opportunity to engage even more, grow, and succeed. It wouldn't be possible to get here without the synergy of team members’ hard work and authors and editors who devote time and their expertise into Open Access book publishing with us.
\n\nOver these years, we have gone from pioneering the scientific Open Access book publishing field to being the world’s largest Open Access book publisher. Nonetheless, our vision has remained the same: to meet the challenges of making relevant knowledge available to the worldwide community under the Open Access model.
\n\nWe are excited about the present, and we look forward to sharing many more successes in the future.
\n\nThank you all for being part of the journey. 5,000 times thank you!
\n\nNow with 5,000 titles available Open Access, which one will you read next?
\n\nRead, share and download for free: https://www.intechopen.com/books
\n\n\n\n
\n'}],latestNews:[{slug:"intechopen-authors-included-in-the-highly-cited-researchers-list-for-2020-20210121",title:"IntechOpen Authors Included in the Highly Cited Researchers List for 2020"},{slug:"intechopen-maintains-position-as-the-world-s-largest-oa-book-publisher-20201218",title:"IntechOpen Maintains Position as the World’s Largest OA Book Publisher"},{slug:"all-intechopen-books-available-on-perlego-20201215",title:"All IntechOpen Books Available on Perlego"},{slug:"oiv-awards-recognizes-intechopen-s-editors-20201127",title:"OIV Awards Recognizes IntechOpen's Editors"},{slug:"intechopen-joins-crossref-s-initiative-for-open-abstracts-i4oa-to-boost-the-discovery-of-research-20201005",title:"IntechOpen joins Crossref's Initiative for Open Abstracts (I4OA) to Boost the Discovery of Research"},{slug:"intechopen-hits-milestone-5-000-open-access-books-published-20200908",title:"IntechOpen hits milestone: 5,000 Open Access books published!"},{slug:"intechopen-books-hosted-on-the-mathworks-book-program-20200819",title:"IntechOpen Books Hosted on the MathWorks Book Program"},{slug:"intechopen-s-chapter-awarded-the-guenther-von-pannewitz-preis-2020-20200715",title:"IntechOpen's Chapter Awarded the Günther-von-Pannewitz-Preis 2020"}]},book:{item:{type:"book",id:"3401",leadTitle:null,fullTitle:"Advances in the Scientific Evaluation of Bladder Cancer and Molecular Basis for Diagnosis and Treatment",title:"Advances in the Scientific Evaluation of Bladder Cancer and Molecular Basis for Diagnosis and Treatment",subtitle:null,reviewType:"peer-reviewed",abstract:"Bladder cancer is the sixth most common cancer in the world affecting more than 300,000 men and women worldwide. 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Enteropathogenic E. coli (EPEC) was the first pathotype of E. coli to be associated with human disease and is a major cause of acute and chronic diarrhea in infants [1, 2]. The low microbial density of the small bowel caused by the forceful peristalsis in this part of the intestine is overcome by EPEC, which can successfully colonize the small intestine of humans [3, 4]. EPEC primarily affects children younger than 2 years old; however some outbreaks of EPEC infection in healthy adults have been associated with large inoculum ingestion [5]. The mechanism of transmission of EPEC is the fecal-oral route. In the 1940s and 1950s, EPEC was an important cause of diarrhea in developed countries with a mortality of 50% during outbreaks, but nowadays the infection by EPEC in industrial countries has a limited importance. In contrast, in low-income countries, EPEC is still an major cause of infant diarrhea [5, 6].
The phenotype that defines EPEC infection is the attaching and effacing (A/E) lesion [2, 7]. By adhering to intestinal epithelial cells, EPEC subverts cytoskeletal processes of the host cell and produces the histopathological feature of the A/E lesion. This lesion, which was first described in 1980 [8], is characterized by the intimate attachment of the bacteria to the intestinal epithelial cells and elongation and effacement of the brush border microvilli. Later on it was shown that infection is also associated with cytoskeletal rearrangements, including the accumulation of polymerized F-actin in pedestal-like structures underneath the attached bacteria [9] (Figure 1). EPEC together with enterohemorrhagic E. coli (EHEC) and Citrobacter rodentium (CR) is a member of the A/E family of bacterial pathogens that colonize the gastrointestinal tract via the A/E lesion. EPEC and EHEC are important human pathogens, while CR is a mouse-restricted pathogen [10, 11, 12, 13].
Localized adherence (LA), intimate attachment, and EPEC A/E lesion formation in the intestinal epithelial surface. At an early stage, EPEC interacts in a non-intimate manner with the intestinal surface mainly through the BFP and EspA filament. After assembly of the translocation pore, EPEC injects translocated intimin receptor (Tir). Ser/Thr phosphorylation of Tir induces its anchoring in the enterocyte plasma membrane, leaving the TirM region exposed for the interaction with intimin. Subsequent Tir-intimin interaction triggers actin polymerization and pedestal formation underneath the attached bacterium. Tir phosphorylation of residue Y474 engages the host adaptor NcK, which later recruits N-WASP and WIP. N-WASP recruits the ARP2/3 complex, which induces actin nucleation and polymerization.
The ability of EPEC to induce A/E lesions is related to a pathogenicity island (PAI) of 35 kb called the locus of enterocyte effacement (LEE) [14]. The LEE comprises 41 genes organized in 5 principal operons (LEE1-LEE5) and several smaller transcriptional units (Figure 2) [15, 16]. Orthologues of LEE are also found in other members of A/E pathogens [11]. The LEE encodes all the structural proteins necessary for the assembly of a filamentous type III secretion system (T3SS) injectisome on the bacterial cell envelope [17, 18]. The LEE also encodes transcriptional regulators (Ler, GrlR, and GrlA), translocator proteins (EspA, EspB, and EspD), six secreted effector proteins (including the translocated intimin receptor), the outer membrane protein intimin, molecular chaperones, and a lytic transglycosylase (EtgA) [19]. The mechanism of LEE regulation is complex and depends on environmental conditions, quorum sensing (QS), and several transcriptional regulators encoded within and outside the LEE [20, 21].
The type III secretion system is a macromolecular transport apparatus that is used by many gram-negative bacterial pathogens (e.g., Shigella, Yersinia, Salmonella) to translocate virulence proteins, called effectors, into the cytosol of infected cells, thereby subverting host cellular functions for the benefit of the pathogen [22]. Since pathogens use this transport apparatus to inject proteins into the host cells, this structure is also known as the injectisome. The EPEC T3SS mediates the translocation of multiple effector proteins during infection. Some of them are encoded in the LEE, whereas others are encoded outside of the LEE being generally referred to as non-LEE effectors (Nle) [23, 24]. EspA filaments link the tip of the injectisome in the bacterial cell wall to a 3–5 nm translocation pore, formed in the plasma membrane of infected cells by the translocator proteins EspB and EspD (Figure 1) [25, 26].
Effectors of EPEC E2348/69. (A) Representation of the LEE island and effector genes espG, espZ, espH, map, tir, and espF. (B) Non-LEE effectors located outside the LEE are localized in integrative elements (IEs) and prophages (PPs). Effector genes are labeled in red. Pseudogenes are specified with asterisk. Scale of 5 kb is indicated at the bottom. Figure from [32].
Typical EPEC is endowed with a plasmid called pMAR2 which contains a 14-gene operon encoding the type IV pilus BFP [27, 28]. The BFP is a rope-like bundle, which allows EPEC to form microcolonies in a pattern called localized adherence and also mediates the initial interaction of bacteria with host cell surfaces (Figure 1) [29, 30, 31].
EPEC tightly regulates its virulence genes in response to environmental conditions such as temperature [16], the increase of the pH of the small intestine [33, 34], and some hormones which are released during stress conditions [20]. Upon EPEC interaction with enterocytes, EspB and EspD proteins are inserted into the host cell membrane and assemble to form a translocation pore [25, 26]. EPEC then injects its own receptor called Tir, which is integrated into the plasma membrane in a hairpin loop topology, with the loop facing the outside of the cell where it serves as a receptor for the bacterial adhesin intimin [35, 36, 37]. Tir-intimin interaction induces clustering and dimerization of Tir, and this activates a signal cascade that starts with the phosphorylation of serine/threonine residues and leads to actin polymerization and pedestal formation underneath the attached bacterium [10, 38]. The most critical event for actin polymerization is the phosphorylation of the cytoplasmic Tir residue Y474 [39]. This induces a signal cascade which recruits the host cell adaptor Nck and N-WASP required to engage and activate the actin-nucleating ARP2/3 complex, which produces the actin nucleation and polymerization. Actin polymerization drives membrane protrusion and pedestal formation [10, 40] (Figure 1). Through the T3SS injectisome, EPEC translocates LEE-encoded effector proteins and additional effectors localized in mobile genetic element outside the LEE (Nle).
Six effector proteins (EspG, EspZ, EspH, Map, Tir, and EspF) are encoded in the LEE island (Figure 2). Most of these, except EspZ, have important functions destabilizing the physiology of the intestinal epithelium, triggering cytoskeleton reorganization, inducing cytotoxicity and electrolyte imbalance which lead to diarrhea [11, 41]. The rapid onset of EPEC-induced diarrhea is likely induced by the cooperative action of Tir, Map, and EspF, which inhibits the sodium-D-glucose transporter (SGLT-1), the major water pump of the small intestine responsible for about 70% of the total fluid uptake [42]. In addition, Map and EspF reduce Na + absorption by the sodium-hydrogen exchanger (NHE3) [43], and EspG1/2 proteins alter the membrane targeting of the Cl-/OH-exchanger (DRA), resulting in reduced Cl-uptake. These processes result in the accumulation of salts in the gut lumen, which drives water loss from the mucosa [44].
Inhibition of endosomal trafficking by EspG1/2 reduces the level of cell surface receptors [45]. In addition, EspF and EspG induce mislocalization of aquaporins (AQP), thereby reducing epithelial water absorption [46]. Furthermore, EspB, Tir, EspF, and Map induce microvillus effacement, and this reduction of absorptive surface likely exacerbates EPEC diarrhea [47]. While EspF and Map synergistically disrupt TJs [48], EspG1/2 induces microtubule disruption contributing to TJ disruption [49]. The effector protein NleA also disrupts TJs by blocking the delivery of new TJ proteins [49, 50, 51]. The disruption of TJs increases intestinal permeability and thereby likely contributes to EPEC-induced diarrhea [52] (Figure 3).
EPEC effector proteins altering epithelial cell function and inducing water loss and diarrhea. Tir, map, and EspF inhibit the sodium-D-glucose transporter. EspF reduces expression of the sodium-hydrogen exchanger NHE3. EspG and EspF induce mislocalization of aquaporins (AqP). EspG1/EspG2 alters membrane targeting of the Cl-/OH-exchanger. EspF, map, NleA, EspG1, and EspG2 disrupt tight junction complexes (TJ). EspB, Tir, EspF, and map induce microvilli effacement.
In EPEC prototype strain E2348/69, 17 functional Nle effectors are encoded in different integrative elements and prophages, frequently associated in gene clusters, with some effectors having duplicated gene copies and/or paralogs in different clusters [53] (Figure 2). EPEC infection is characterized by a weak inflammatory response [54]. Previous studies have shown that most Nle effectors and some LEE effectors inhibit the host immune response, which favors bacterial survival (Figure 4). Although NleF and NleH2 activate the NF-κB inflammatory pathway during early infection (ref), EPEC translocates several effectors that dampen the proinflammatory pathways of the cell [11]. Thus, a large number of Nle effectors inhibit host inflammation by different mechanisms, such as inhibition of the NF-κB (NleB, C, E, and H) and MAPK proinflammatory pathways (NleC and D) [55, 56, 57, 58], inhibition of the canonical (NleA) and noncanonical (NleF) inflammasomes [59], and inhibition of proliferation of lymphocytes and interleukin production (LifA) [60, 61].
Schematic representation of multifunctional and overlapping effectors to control host immune response. The NF-κB proinflammatory pathway is activated by NleF and NleH2 and is inhibited by NleE, NleB, NleH1, Tir, and NleC. NleC and NleD inhibit the MAPK proinflammatory pathway. EspF, EspJ, EspH, and EspB prevent macrophage phagocytosis. NleA disrupts inflammasome activation, and LifA inhibits IL-2 and IL-4 production and lymphocyte proliferation. While EspF and map induce intrinsic apoptosis, EspZ counteracts these effects by stabilizing mitochondrial membrane potential. NleH1/NleH2 and NleF inhibit intrinsic apoptosis, and NleF, NleD, and NleB counteract extrinsic apoptosis.
The control of the epithelial cell death response to microbial infection is pivotal for pathogens and the host. Pathogens that are colonizing the epithelium need to prevent cell death to preserve their replicative foothold; by contrast, the host needs to eliminate infected cells in order to minimize tissue damage [62]. During infection of the intestinal epithelial cells, surface properties of EPEC are recognized by cell surface death receptors and induce extrinsic apoptotic pathways, while T3SS effectors (Map and EspF) trigger cytochrome c release, activation of caspases, and downstream intrinsic apoptotic pathways [11, 24]. Interestingly, early stages of apoptosis can be observed during EPEC infection, but late stages are not evident because EPEC translocates effector proteins that antagonize these pro-apoptotic effects. NleD and NleB interfere with the pro-apoptotic death receptor signaling and disrupt the downstream extrinsic apoptosis [63, 64]. NLeH1/2 and EspZ also inhibit intrinsic apoptosis and promote host cell survival [65, 66, 67] (Figure 4). NleF directly inhibits caspases involved in both intrinsic and extrinsic apoptosis pathways, including caspases 4, 8, and 9 [68]. In addition, EspZ localizes to the cytoplasmic side of the plasma membrane at the site of bacterial attachment and interacts with the translocator protein EspD. It has been proposed that EspZ indirectly prevents cell death by downregulating protein translocation and protecting cells from an overdose of effector proteins. Consistently, a ΔespZ mutant was found to be highly cytotoxic [69]. EPEC effectors are injected in a regulated manner to guarantee the success of infection. While the pro-survival effector EspZ is translocated at the early stages of infection, the pro-apoptotic effectors EspF and Map follow later [70].
Most research on EPEC effectors has been conducted by generating deletion mutants in a single or a few effector genes that are later complemented with multicopy plasmids overexpressing the effector(s). In addition, ectopic expression of individual effectors by plasmid transfection of the host cell has been applied. Both situations are prone to effector overexpression resulting in nonphysiological levels of effectors inside the host cell, which could alter effector activities. In addition, effectors often have synergistic and overlapping functions that cannot be fully appreciated by single mutations and individual transfection experiments [11, 54]. In order to overcome these limitations, we employed a marker-less gene deletion strategy to delete the whole repertoire of known effector genes found in the genome of the prototypical EPEC strain E2348/69 [32]. The genome engineering method for sequential deletion of EPEC effectors was based on the marker-less gene deletion technique described by Posfai et al. [71] and is illustrated in Figure 5.
Marker-less gene deletion strategy of EPEC effector genes. Deletions using pGE-suicide plasmids with I-SceI sites and mutant alleles assembled by fusing homology regions (HRs) flanking the targeted effector gene(s). Co-integrants are identified by the Kanamycin resistance phenotype. Expression of the I-SceI in vivo from helper plasmid induces double-strand brakes that are repaired by homologous recombination. Depending on the HRs involved in this second recombination, either the WT or the mutant allele can be obtained. Figure from [32].
Using this strategy, a set of EPEC mutants with sequential deletions of effectors was generated (Table 1), ultimately resulting in strains expressing only Tir and EspZ (EPEC2), Tir (EPEC1), and the effector-less strain EPEC0 (Table 1). This approach proved to be effective to specifically modify the genome of EPEC E2348/69, avoiding the introduction of unintended alterations in the genome and leaving no sequence “scars” or antibiotic resistance genes in the chromosome as demonstrated by whole-genome sequencing [32]. Besides, the deletion mutant strains showed normal growth and maintained functional T3SS injectisomes. In addition, they allowed the translocation of individual effectors from single-copy chromosomal genes under endogenous regulation, showing the expected phenotypes without the background of the other effectors [32]. Hence these mutant strains are an excellent tool to investigate the role of individual effectors and specific combinations maintaining physiological protein levels in the context of infection.
Strain | Effector genes remaining* |
---|---|
WT | All |
EPEC2 | espZ and tir |
EPEC1 | tir |
EPEC0 | None |
EPEC9 | espZ + tir + IE2 + IE5 + IE6 + PP2 + PP3 + PP4 + PP6 |
EPEC2-LEE+ | espZ + tir + map + espH + espF + espG |
EPEC mutant strains generated with the marker-less deletion strategy.
Encoded effectors in the indicated IEs and PPs.
When EPEC bacteria adhere in vitro to cultured cells, there is an accumulation of actin filaments in the cytoplasm beneath the adherent bacteria, due to a signal cascade triggered by intimin-Tir interaction [35, 38]. Using the effector deletion mutants of EPEC, we demonstrated that the LEE effector Tir along with intimin is necessary and sufficient to induce these cytoskeletal rearrangements during in vitro infection of HeLa cells. Strains EPEC2 (bearing EspZ and Tir) and EPEC1 (bearing only Tir) were able to induce actin-pedestal formation underneath attached bacteria similar to the EPEC wild type (WT) (Figure 6). As expected because of the essential role of Tir in this process, infection of HeLa cells with the effector-less mutant EPEC0 did not induce any actin-pedestal formation (Figure 6). These data demonstrate that the individual translocation of Tir by EPEC1 is sufficient to trigger actin pedestals in HeLa cells and that non-LEE effectors are dispensable for this phenotype during in vitro infection of cultured cells.
Infection of HeLa cells with EPEC WT and effector mutant strains. Immunofluorescence confocal microscopy of HeLa cells infected with EPEC WT, EPEC2, EPEC1, and EPEC0 for 1.5 h using a MOI of 200. EPEC is labeled with anti-intimin-280 serum (green), actin is stained with TRITC phalloidin (red), and cell nuclei are labeled with DAPI (gray). Actin polymerization beneath adherent bacteria is observed in EPEC WT, EPEC2, and EPEC1. Scale bar 5 μm. Figure from [32].
EPEC pathogenic mechanisms have been widely investigated by in vitro infection of cultured epithelial cell lines, albeit in most cases these cells are non-polarized and are not from intestinal origin. In addition, EPEC infection studies in vivo are hindered because EPEC is a human-restricted pathogen [72]. A surrogate model established to investigate A/E pathogenesis in vivo is the mouse pathogen Citrobacter rodentium (CR) [12, 13]. Although Citrobacter infection in vitro requires Tir phosphorylation for actin-pedestal formation in cell lines, Tir phosphorylation-deficient mutants still colonize the mouse gut and induce A/E lesion formation and crypt hyperplasia typical of CR infection [73]. This result highlights the necessity of a model for EPEC infection closer to the in vivo conditions in the human gut. A good established model to study EPEC-host interactions is the infection of in vitro cultured human intestinal biopsies, which allows the formation of A/E lesions undistinguishable from those observed in vivo in biopsies of patients with EPEC-induced diarrhea [4, 36, 74]. Similar to results obtained in CR-infected mice, Tir phosphorylation was not necessary for EPEC A/E lesion formation in human intestinal biopsies [75]. Surprisingly, when EPEC2 and EPEC1 deletion mutants were used to infect human duodenal biopsies, none of the infected biopsies showed A/E lesions (Table 2 and Figure 7), which contrasts with the pedestal formation observed in HeLa cells. Thus, intimin and Tir are not sufficient to induce A/E lesions in the intestinal tissue, and the IVOC model was used to identify additional LEE or non-LEE effector(s) required for A/E lesion formation. For this purpose, two additional effector mutant strains were tested: EPEC2-LEE+ (carrying all LEE effectors) and EPEC9 (carrying EspZ, Tir, and all non-LEE effectors). Whereas infection with EPEC2-LEE+ did not reveal A/E lesions, infection with EPEC9 induced A/E lesions to a similar level as the wild-type strain (Table 2 and Figure 7). It was previously reported that the LEE island is sufficient to confer the A/E phenotype to E. coli K-12 in the infection of colon carcinoma cell lines [76]. However, our results indicate that the LEE is not sufficient for A/E lesion formation in human mucosal tissue and that non-LEE effectors are required [32].
Strain | Effector genes remaining* | Biopsies with A/E lesions positive/total (%) |
---|---|---|
WT | All | 13/17 (76) |
EPEC2 | espZ and tir | 0/6 (0) |
EPEC1 | tir | 0/6 (0) |
EPEC0 | None | 0/5 (0) |
EPEC9 | espZ + tir + IE2 + IE5 + IE6 + PP2 + PP3 + PP4 + PP6 | 5/6 (83) |
EPEC2-LEE+ | espZ + tir + map + espH + espF + espG | 0/5 (0) |
Human duodenal biopsies infected by EPEC WT and EPEC effector mutants.
Encoded effectors in the indicated IEs and PPs.
Scanning electron microscopy of human duodenal biopsies infected with EPEC WT and mutant strains EPEC2, EPEC1, EPEC0, EPEC9, and EPEC2-LEE+. EPEC WT and EPEC9 induce characteristic A/E lesions with bacterial microcolony formation (asterisk) and microvilli elongation around bacterial colonies (arrowheads). In contrast, biopsies infected with EPEC2, EPEC1, EPEC0, and EPEC2-LEE+ lack adherent bacteria and A/E lesions and show a normal microvillous brush border. Scale bar 2 μm. Figure from [32].
The marker-less gene deletion strategy enabled the generation of effector-less strains of EPEC O127:H6 using the prototypical strain E2348/69 [32]. Given the conservation of the recombination machinery among E. coli strains, it is likely that this strategy could be applied to other A/E pathogens, E. coli pathogens, and other bacteria. The effector mutant strains can be useful to study the role of individual effectors and of combinations of effectors in pathogenesis. An individual effector or a defined combination can be inserted in the effector-less strains in their endogenous genomic loci to obtain physiological expression levels and regulation. In cell culture infections, all EPEC effector mutant strains carrying intimin and Tir were able to trigger actin-rich pedestal-like structures underneath attached bacteria. On the other hand, when the infection was performed in human intestinal tissues, translocation of Tir alone was insufficient to induce A/E lesions. Furthermore, an EPEC deletion mutant maintaining all LEE effectors and devoid of all non-LEE effectors (EPEC2-LEE+) was still unable to induce A/E lesions in human intestinal biopsies. In contrast, an EPEC strain producing the complete repertoire of non-LEE effectors and devoid of LEE effectors, except Tir and EspZ, formed A/E lesions in intestinal tissue at wild-type levels [32]. Thus, these experiments revealed that non-LEE effectors are needed for A/E lesion formation in human intestinal tissue.
In addition to their potential for basic studies, the EPEC effector mutant strains may have different applications. For instance, EPEC (and other pathogenic) strains lacking multiple effectors are likely to be strongly attenuated, but they maintain the external antigenicity of the wild-type strain. Thus, an EPEC mutant strain with a functional T3SS and the minimum set of effectors necessary to colonize the intestinal surface could be a good vaccine candidate. Further, EPEC mutant strains with the ability to attach to the human intestine could also be engineered to translocate heterologous protein antigens to generate protection against other enteric pathogens causing diarrhea, including EHEC strains [77, 78, 79, 80]. Lastly, the EPEC effector mutant strains may also have the potential to deliver therapeutic proteins to the intestinal epithelium, for instance, to combat inflammation and autoimmune disorders in the gastrointestinal tract [81].
We acknowledge support for the publication fee by the CSIC Open Access Publication Support Initiative through its Unit of Information Resources for Research (URICI).
The Internet has irrevocably changed the dynamics of scholarly communication and publishing. Consequently, we find it necessary to indicate, unambiguously, our definition of what we consider to be a published scientific work.
",metaTitle:"Prior Publication Policy",metaDescription:"Prior Publication Policy",metaKeywords:null,canonicalURL:"/page/prior-publication-policy",contentRaw:'[{"type":"htmlEditorComponent","content":"A significant number of working papers, early drafts, and similar work in progress are openly shared online between members of the scientific community. It has become common to announce one’s own research on a personal website or a blog to gather comments and suggestions from other researchers. Such works and online postings are, indeed, published in the sense that they are made publicly available. However, this does not mean that if submitted for publication by IntechOpen they are not original works. We differentiate between reviewed and non-reviewed works when determining whether a work is original and has been published in a scholarly sense or not.
\\n\\nThe significance of Peer Review cannot be overstated when it comes to defining, in our terms, what constitutes a published scientific work. Peer Review is widely considered to be the cornerstone of modern publishing processes and the key value-adding contribution to a scholarly manuscript that a publisher can make.
\\n\\nOther than the issue of originality, research misconduct is another major issue that all publishers have to address. IntechOpen’s Retraction & Correction Policy and various publication ethics guidelines identify both redundant publication and (self)plagiarism to fall within the definition of research misconduct, thus constituting grounds for rejection or the issue of a Retraction if the work has already been published.
\\n\\nIn order to facilitate the tracking of a manuscript’s publishing history and its development from its earliest draft to the manuscript submitted, we encourage Authors to disclose any instances of a manuscript’s prior publication, whether it be through a conference presentation, a newspaper article, a working paper publicly available in a repository or a blog post.
\\n\\nA note to the Academic Editor containing detailed information about a submitted manuscript’s previous public availability is the preferred means of reporting prior publication. This helps us determine if there are any earlier versions of a manuscript that should be disclosed to our readers or if any of those earlier versions should be cited and listed in a manuscript’s references.
\\n\\nSome basic information about the editorial treatment of different varieties of prior publication is laid out below:
\\n\\n1. CONFERENCE PAPERS & PRESENTATIONS
\\n\\nGiven that conference papers and presentations generally pass through some sort of peer or editorial review, we consider them to be published in the accepted scholarly sense, particularly if they are published as a part of conference proceedings.
\\n\\nAll submitted manuscripts originating from a previously published conference paper must contain at least 50% of new original content to be accepted for review and considered for publication.
\\n\\nAuthors are required to report any links their manuscript might have with their earlier conference papers and presentations in a note to the Academic Editor, as well as in the manuscript itself. Additionally, Authors should obtain any necessary permissions from the publisher of their conference paper if copyright transfer occurred during the publishing process. Failure to do so may prevent Us from publishing an otherwise worthy work.
\\n\\n2. NEWSPAPER & MAGAZINE ARTICLES
\\n\\nNewspaper and magazine articles usually do not pass through any extensive peer or editorial review and we do not consider them to be published in the scholarly sense. Articles appearing in newspapers and magazines rarely possess the depth and structure characteristic of scholarly articles.
\\n\\nSubmitted manuscripts stemming from a previous newspaper or magazine article will be accepted for review and considered for publication. However, Authors are strongly advised to report any such publication in an accompanying note to the External Editor.
\\n\\nAs with the conference papers and presentations, Authors should obtain any necessary permissions from the newspaper or magazine that published the work, and indicate that they have done so in a note to the External Editor.
\\n\\n3. GREY LITERATURE
\\n\\nWhite papers, working papers, technical reports and all other forms of papers which fall within the scope of the ‘Luxembourg definition’ of grey literature do not pass through any extensive peer or editorial review and we do not consider them to be published in the scholarly sense.
\\n\\nAlthough such papers are regularly made publicly available via personal websites and institutional repositories, their general purpose is to gather comments and feedback from Authors’ colleagues in order to further improve a manuscript intended for future publication.
\\n\\nWhen submitting their work, Authors are required to disclose the existence of any publicly available earlier drafts in a note to the Academic Editor. In cases where earlier drafts of the submitted version of the manuscript are publicly available, any overlap between the versions will generally not be considered an instance of self-plagiarism.
\\n\\n4. SOCIAL MEDIA, BLOG & MESSAGE BOARD POSTINGS
\\n\\nWe feel that social media, blogs and message boards are generally used with the same intention as grey literature, to formulate ideas for a manuscript and gather early feedback from like-minded researchers in order to improve a particular piece of work before submitting it for publication. Therefore, we do not consider such internet postings to be publication in the scholarly sense.
\\n\\nNevertheless, Authors are encouraged to disclose the existence of any internet postings in which they outline and describe their research or posted passages of their manuscripts in a note to the Academic Editor. Please note that we will not strictly enforce this request in the same way that we would instructions we consider to be part of our conditions of acceptance for publication. We understand that it may be difficult to keep track of all one’s internet postings in which the researcher´s current work might be mentioned.
\\n\\nIn cases where there is any overlap between the Author´s submitted manuscript and related internet postings, we will generally not consider it to be an instance of self-plagiarism. This also holds true for any co-Author as well.
\\n\\nFor more information on this policy please contact permissions@intechopen.com.
\\n\\nPolicy last updated: 2017-03-20
\\n"}]'},components:[{type:"htmlEditorComponent",content:'A significant number of working papers, early drafts, and similar work in progress are openly shared online between members of the scientific community. It has become common to announce one’s own research on a personal website or a blog to gather comments and suggestions from other researchers. Such works and online postings are, indeed, published in the sense that they are made publicly available. However, this does not mean that if submitted for publication by IntechOpen they are not original works. We differentiate between reviewed and non-reviewed works when determining whether a work is original and has been published in a scholarly sense or not.
\n\nThe significance of Peer Review cannot be overstated when it comes to defining, in our terms, what constitutes a published scientific work. Peer Review is widely considered to be the cornerstone of modern publishing processes and the key value-adding contribution to a scholarly manuscript that a publisher can make.
\n\nOther than the issue of originality, research misconduct is another major issue that all publishers have to address. IntechOpen’s Retraction & Correction Policy and various publication ethics guidelines identify both redundant publication and (self)plagiarism to fall within the definition of research misconduct, thus constituting grounds for rejection or the issue of a Retraction if the work has already been published.
\n\nIn order to facilitate the tracking of a manuscript’s publishing history and its development from its earliest draft to the manuscript submitted, we encourage Authors to disclose any instances of a manuscript’s prior publication, whether it be through a conference presentation, a newspaper article, a working paper publicly available in a repository or a blog post.
\n\nA note to the Academic Editor containing detailed information about a submitted manuscript’s previous public availability is the preferred means of reporting prior publication. This helps us determine if there are any earlier versions of a manuscript that should be disclosed to our readers or if any of those earlier versions should be cited and listed in a manuscript’s references.
\n\nSome basic information about the editorial treatment of different varieties of prior publication is laid out below:
\n\n1. CONFERENCE PAPERS & PRESENTATIONS
\n\nGiven that conference papers and presentations generally pass through some sort of peer or editorial review, we consider them to be published in the accepted scholarly sense, particularly if they are published as a part of conference proceedings.
\n\nAll submitted manuscripts originating from a previously published conference paper must contain at least 50% of new original content to be accepted for review and considered for publication.
\n\nAuthors are required to report any links their manuscript might have with their earlier conference papers and presentations in a note to the Academic Editor, as well as in the manuscript itself. Additionally, Authors should obtain any necessary permissions from the publisher of their conference paper if copyright transfer occurred during the publishing process. Failure to do so may prevent Us from publishing an otherwise worthy work.
\n\n2. NEWSPAPER & MAGAZINE ARTICLES
\n\nNewspaper and magazine articles usually do not pass through any extensive peer or editorial review and we do not consider them to be published in the scholarly sense. Articles appearing in newspapers and magazines rarely possess the depth and structure characteristic of scholarly articles.
\n\nSubmitted manuscripts stemming from a previous newspaper or magazine article will be accepted for review and considered for publication. However, Authors are strongly advised to report any such publication in an accompanying note to the External Editor.
\n\nAs with the conference papers and presentations, Authors should obtain any necessary permissions from the newspaper or magazine that published the work, and indicate that they have done so in a note to the External Editor.
\n\n3. GREY LITERATURE
\n\nWhite papers, working papers, technical reports and all other forms of papers which fall within the scope of the ‘Luxembourg definition’ of grey literature do not pass through any extensive peer or editorial review and we do not consider them to be published in the scholarly sense.
\n\nAlthough such papers are regularly made publicly available via personal websites and institutional repositories, their general purpose is to gather comments and feedback from Authors’ colleagues in order to further improve a manuscript intended for future publication.
\n\nWhen submitting their work, Authors are required to disclose the existence of any publicly available earlier drafts in a note to the Academic Editor. In cases where earlier drafts of the submitted version of the manuscript are publicly available, any overlap between the versions will generally not be considered an instance of self-plagiarism.
\n\n4. SOCIAL MEDIA, BLOG & MESSAGE BOARD POSTINGS
\n\nWe feel that social media, blogs and message boards are generally used with the same intention as grey literature, to formulate ideas for a manuscript and gather early feedback from like-minded researchers in order to improve a particular piece of work before submitting it for publication. Therefore, we do not consider such internet postings to be publication in the scholarly sense.
\n\nNevertheless, Authors are encouraged to disclose the existence of any internet postings in which they outline and describe their research or posted passages of their manuscripts in a note to the Academic Editor. Please note that we will not strictly enforce this request in the same way that we would instructions we consider to be part of our conditions of acceptance for publication. We understand that it may be difficult to keep track of all one’s internet postings in which the researcher´s current work might be mentioned.
\n\nIn cases where there is any overlap between the Author´s submitted manuscript and related internet postings, we will generally not consider it to be an instance of self-plagiarism. This also holds true for any co-Author as well.
\n\nFor more information on this policy please contact permissions@intechopen.com.
\n\nPolicy last updated: 2017-03-20
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