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n2:pmcid: PMC0
bibo:doi: 10.1084%2Fjem.20071022
n4:contains: _:vb3044267 _:vb3044251 _:vb3044243

Subject Item: _:vb3044243
rdf:type: n4:Section
dc:title: materials and methods
n4:contains: _:vb3044248 _:vb3044249 _:vb3044250 _:vb3044244 _:vb3044245 _:vb3044246 _:vb3044247

Subject Item: _:vb3044244
rdf:type: n2:Context
rdf:value: Primary IECs were isolated from the intestinal tissue at different ages based on a published protocol (>>10<<). All tissue, except that from 0 h mice, was obtained from spontaneously delivered newborns. 0-h-old IECs were isolated from mice born by caesarean section after natural birth of the first sibling to ensure maturity of the obtained
n2:mentions: n3:16606665

Subject Item: _:vb3044245
rdf:type: n2:Context
rdf:value: Cryptin 4(forward [Defcrp130], 5′-AAGAGACTAAAACTGAGGAGCAGC-3′; reverse, 5′-CGGCGGGGGCAGCAGTA-3′), and Cryptin 5 (forward [Defcrp130], 5′-AAGAGACTAAAACTGAGGAGCAGC-3′; reverse, 5′-GCAGCAGAATACGAAAGT-3′) were as previously described (>>9<<). Hprt1 (forward, 5′-TGATCAGTCAACGGGGGACA-3′; reverse, 5′-TTCGAGAGGTCCTTTTCACCA-3′) and β2-microglobulin (b2M; forward, 5′-TGGTGCTTGTCTCACTGAML-3′; reverse, 5′-CCGTTCTTCAGCATTTGGAT-3′) were used as endogenous controls to normalize gene
n2:mentions: n3:9038894

Subject Item: _:vb3044246
rdf:type: n2:Context
rdf:value: Cationic antimicrobial peptides were extracted from isolated IECs of newborn and adult C57BL/6 mice or m-ICcl2 cell lysate, according to a published protocol (>>12<<). The antibacterial activity of HPLC fractions was analyzed using a zone inhibition assay using B. megaterium strain 11 (11).
n2:mentions: n3:15235601

Subject Item: _:vb3044247
rdf:type: n2:Context
rdf:value: The antibacterial activity of HPLC fractions was analyzed using a zone inhibition assay using B. megaterium strain 11 (>>11<<). Cell-culture supernatant was purified using Sep-Pak Light tC18 cartridges (Waters Corp.) before antibacterial testing and mass spectrometric analysis. Matrix-assisted laser desorption-ionization time-of-flight mass spectrometry analysis
n2:mentions: n3:11010975

Subject Item: _:vb3044248
rdf:type: n2:Context
rdf:value: Matrix-assisted laser desorption-ionization time-of-flight mass spectrometry analysis was performed with a Reflex III instrument (Bruker Daltronics), as recently described (>>11<<).
n2:mentions: n3:11010975

Subject Item: _:vb3044249
rdf:type: n2:Context
rdf:value: Immunoblotting was performed as recently described (>>10<<). Membranes were incubated overnight at 4°C, with the primary antibody provided by B.
n2:mentions: n3:16606665

Subject Item: _:vb3044250
rdf:type: n2:Context
rdf:value: Typhimurium, as previously described (>>37<<). 104 bacteria grown to mid-logarithmic phase were coincubated in 100 μl with or without the addition of peptide at the concentrations indicated in the figures (0.5, 1, 5, 10, 20, and 50 μg/ml).
n2:mentions: n3:6628360

Subject Item: _:vb3044251
rdf:type: n4:Section
dc:title: results
n4:contains: _:vb3044252 _:vb3044253 _:vb3044254 _:vb3044255 _:vb3044264 _:vb3044265 _:vb3044266 _:vb3044256 _:vb3044257 _:vb3044258 _:vb3044259 _:vb3044260 _:vb3044261 _:vb3044262 _:vb3044263

Subject Item: _:vb3044252
rdf:type: n2:Context
rdf:value: investigate gene expression of antimicrobial peptides during postnatal intestinal development, we quantitatively examined messenger RNA (mRNA) levels in highly enriched primary IECs prepared according to a recently established protocol (>>10<<). Total RNA of IECs from mouse small intestinal tissue was isolated at the time of birth or at the ages of 6 and 24 h, or 3, 6, 14, 21, and 28 d and analyzed by real-time PCR.
n2:mentions: n3:16606665

Subject Item: _:vb3044253
rdf:type: n2:Context
rdf:value: All fractions were analyzed for antimicrobial activity against Bacillus megaterium (strain Bm11) using a standardized agar diffusion assay (>>11<<). In addition, mass spectrometry was performed on positive fractions to identify antimicrobially active substances. The presence of a large variety of cryptdins and CRS peptide homo- and heterodimers was confirmed by mass spectrometric
n2:mentions: n3:11010975

Subject Item: _:vb3044254
rdf:type: n2:Context
rdf:value: variety of cryptdins and CRS peptide homo- and heterodimers was confirmed by mass spectrometric analysis in fractions 45–55 from adult IECs (day 28) associated with significant antibacterial activity, as recently reported (Fig. 2 A) (>>11<<, 12).
n2:mentions: n3:11010975

Subject Item: _:vb3044255
rdf:type: n2:Context
rdf:value: 2 A) (11, >>12<<).
n2:mentions: n3:15235601

Subject Item: _:vb3044256
rdf:type: n2:Context
rdf:value: Alternatively, trypsin activity has recently been identified in mouse small intestinal tissue (>>13<<). Thus, constitutive expression and production of the processed mature form of CRAMP was detected in small intestinal epithelium restricted to the neonatal period.
n2:mentions: n3:9736042

Subject Item: _:vb3044257
rdf:type: n2:Context
rdf:value: a highly differentiated and polarized IEC line derived from embryonic day 20 fetuses of transgenic mice carrying the large T antigen of the simian virus 40 under the control of the 5′ regulatory sequence from the L-pyruvate kinase gene (>>14<<). Although none of the established enteric antimicrobial peptides, such as α-defensins, CRS peptides, or angiogenin 4, were detectable by RT-PCR (Fig.
n2:mentions: n3:8764149

Subject Item: _:vb3044258
rdf:type: n2:Context
rdf:value: Cathelicidins have been reported to promote cell differentiation and angiogenesis, and enhance cell proliferation and migration (>>15<<, 16). Neonatal CRAMP expression might thereby contribute to postnatal organ development and intestinal cell differentiation.
n2:mentions: n3:12603850

Subject Item: _:vb3044259
rdf:type: n2:Context
rdf:value: Cathelicidins have been reported to promote cell differentiation and angiogenesis, and enhance cell proliferation and migration (15, >>16<<). Neonatal CRAMP expression might thereby contribute to postnatal organ development and intestinal cell differentiation.
n2:mentions: n3:16177113

Subject Item: _:vb3044260
rdf:type: n2:Context
rdf:value: Although CRAMP significantly inhibited LPS-mediated stimulation of naive epithelial cells similar to other antimicrobial peptides, it showed no effect on endotoxin-tolerant epithelial cells (Fig. S1, D and E) (>>12<<, 17).
n2:mentions: n3:15235601

Subject Item: _:vb3044261
rdf:type: n2:Context
rdf:value: Although CRAMP significantly inhibited LPS-mediated stimulation of naive epithelial cells similar to other antimicrobial peptides, it showed no effect on endotoxin-tolerant epithelial cells (Fig. S1, D and E) (12, >>17<<).
n2:mentions: n3:16456005

Subject Item: _:vb3044262
rdf:type: n2:Context
rdf:value: Mucosal colonization of the intestinal tract starts immediately after birth, leading to an increasingly dense and complex microbial flora during the first weeks of life (>>5<<). To examine a possible effect of microbial colonization on the disappearance of CRAMP expression, conventionally bred and germ-free–bred animals were compared.
n2:mentions: n3:12215177

Subject Item: _:vb3044263
rdf:type: n2:Context
rdf:value: These data are in accordance with a recent report on low Wnt signaling activity in the intervillus region directly after birth and increased activity in the crypt area of adult individuals (>>18<<). They further suggest that CRAMP expression is limited to the neonate epithelium and lost during the developmental changes that occur in the postnatal period associated with increased IEC proliferation and differentiation.
n2:mentions: n3:17681174

Subject Item: _:vb3044264
rdf:type: n2:Context
rdf:value: host protection, we next tested the CRAMP-mediated antibacterial activity against gut commensal bacteria, as well as defined pathogenic bacteria known to be transmitted to the neonate organism during passage through the birth canal (>>6<<). Rapid bacterial killing resulting in a 3–4 log decrease of viable bacteria after 2 h was noted against selected Gram-positive and -negative intestinal commensal bacteria of the small intestine such as S. gallinaceus, Lactobacillus
n2:mentions: n3:14526331

Subject Item: _:vb3044265
rdf:type: n2:Context
rdf:value: Because of structural differences of the epithelial receptor E-cadherin between mice and men, the invasion rate of L. monocytogenes after oral infection of mice is very low and results in only a marginal local inflammatory reaction (>>19<<, 20). This allows application of significant bacterial numbers, minimizes the possible effect of infiltrating granulocytes, and, thus, favors the analysis of the intraluminal intestinal host defense.
n2:mentions: n3:10406800

Subject Item: _:vb3044266
rdf:type: n2:Context
rdf:value: Because of structural differences of the epithelial receptor E-cadherin between mice and men, the invasion rate of L. monocytogenes after oral infection of mice is very low and results in only a marginal local inflammatory reaction (19, >>20<<). This allows application of significant bacterial numbers, minimizes the possible effect of infiltrating granulocytes, and, thus, favors the analysis of the intraluminal intestinal host defense.
n2:mentions: n3:11387478

Subject Item: _:vb3044267
rdf:type: n4:Section
dc:title: discussion
n4:contains: _:vb3044268 _:vb3044269 _:vb3044270 _:vb3044271 _:vb3044280 _:vb3044281 _:vb3044282 _:vb3044283 _:vb3044284 _:vb3044285 _:vb3044286 _:vb3044272 _:vb3044273 _:vb3044274 _:vb3044275 _:vb3044276 _:vb3044277 _:vb3044278 _:vb3044279

Subject Item: _:vb3044268
rdf:type: n2:Context
rdf:value: Importantly, low expression of Paneth cell–derived antimicrobial peptides has also been noted in human neonates (>>21<<). Besides infection via the transplacental route during pregnancy, several microbial organisms are transmitted from the mother to the newborn during passage through the birth canal, gain entrance via the intestinal tract, and may evoke
n2:mentions: n3:8626737

Subject Item: _:vb3044269
rdf:type: n2:Context
rdf:value: Similarly, CRAMP expression has been detected in vernix caseosa and neonate skin (>>22<<–24). Together with our results, these data suggest that CRAMP plays a prominent role in the protection of the newborn.
n2:mentions: n3:15661923 n3:15565791 n3:12612195

Subject Item: _:vb3044270
rdf:type: n2:Context
rdf:value: Expression of the precursor peptide has been noted in myeloid cells, keratinocytes, and epithelial cells from the skin, lung, stomach, colon, meninges, and eye, as well as endothelial cells (>>25<<–28). Although they represent a large and important group of antimicrobial peptides in a variety of mammals, only one member, LL-37 and CRAMP, is encoded in humans and mice, respectively. Significant cathelicidin expression has previously
n2:mentions: n3:14724813 n3:12387964 n3:12692061 n3:11719807

Subject Item: _:vb3044271
rdf:type: n2:Context
rdf:value: cathelicidin expression has previously been noted in the gastrointestinal tract, such as human gastric or colonic tissue, with higher expression levels in individuals with persistent bacterial infection or chronic inflammation (>>27<<, 29). Although our results obtained using highly purified primary IECs indicated the absence of constitutive Cramp expression in adult individuals, Cramp mRNA was recently noted in total adult small intestinal tissue (30).
n2:mentions: n3:14724813

Subject Item: _:vb3044272
rdf:type: n2:Context
rdf:value: cathelicidin expression has previously been noted in the gastrointestinal tract, such as human gastric or colonic tissue, with higher expression levels in individuals with persistent bacterial infection or chronic inflammation (27, >>29<<). Although our results obtained using highly purified primary IECs indicated the absence of constitutive Cramp expression in adult individuals, Cramp mRNA was recently noted in total adult small intestinal tissue (30).
n2:mentions: n3:16702850

Subject Item: _:vb3044273
rdf:type: n2:Context
rdf:value: Although our results obtained using highly purified primary IECs indicated the absence of constitutive Cramp expression in adult individuals, Cramp mRNA was recently noted in total adult small intestinal tissue (>>30<<). The different results might reflect expression by resident tissue myeloid cells. However, we cannot rule out the reappearance of small intestinal epithelial Cramp expression in adult individuals during conditions such as tumorigenesis
n2:mentions: n3:9148921

Subject Item: _:vb3044274
rdf:type: n2:Context
rdf:value: lack of cathelicidin synthesis was associated with clinical disease, such as periodontitis, in human individuals with congenital neutropenia (Kostmann syndrome) and enhanced susceptibility to recurrent urinary tract infection (>>26<<, 31). CRAMP expression also conferred protection from invasive skin infection by group A streptococci and Citrobacter rodentium–induced colitis in mice (25, 32).
n2:mentions: n3:12387964

Subject Item: _:vb3044275
rdf:type: n2:Context
rdf:value: lack of cathelicidin synthesis was associated with clinical disease, such as periodontitis, in human individuals with congenital neutropenia (Kostmann syndrome) and enhanced susceptibility to recurrent urinary tract infection (26, >>31<<). CRAMP expression also conferred protection from invasive skin infection by group A streptococci and Citrobacter rodentium–induced colitis in mice (25, 32).
n2:mentions: n3:16751768

Subject Item: _:vb3044276
rdf:type: n2:Context
rdf:value: CRAMP expression also conferred protection from invasive skin infection by group A streptococci and Citrobacter rodentium–induced colitis in mice (>>25<<, 32). The clinical importance of CRAMP was further underlined by the description of mechanisms to gain resistance against peptide-mediated killing by several human pathogenic bacteria (33, 34). In addition to the broad and potent
n2:mentions: n3:11719807

Subject Item: _:vb3044277
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rdf:value: CRAMP expression also conferred protection from invasive skin infection by group A streptococci and Citrobacter rodentium–induced colitis in mice (25, >>32<<). The clinical importance of CRAMP was further underlined by the description of mechanisms to gain resistance against peptide-mediated killing by several human pathogenic bacteria (33, 34). In addition to the broad and potent
n2:mentions: n3:15814717

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rdf:value: The clinical importance of CRAMP was further underlined by the description of mechanisms to gain resistance against peptide-mediated killing by several human pathogenic bacteria (>>33<<, 34). In addition to the broad and potent antibacterial activity, cathelicidin-derived antimicrobial peptides have also been noted to exert other biological activities so as to promote cell differentiation and to stimulate angiogenesis
n2:mentions: n3:15661016

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rdf:value: The clinical importance of CRAMP was further underlined by the description of mechanisms to gain resistance against peptide-mediated killing by several human pathogenic bacteria (33, >>34<<). In addition to the broad and potent antibacterial activity, cathelicidin-derived antimicrobial peptides have also been noted to exert other biological activities so as to promote cell differentiation and to stimulate angiogenesis and
n2:mentions: n3:12366839

Subject Item: _:vb3044280
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rdf:value: potent antibacterial activity, cathelicidin-derived antimicrobial peptides have also been noted to exert other biological activities so as to promote cell differentiation and to stimulate angiogenesis and epithelial cell proliferation (>>15<<, 16). Although our results do not support an involvement of epithelial CRAMP in the postnatal intestinal organ development, other CRAMP-mediated biological effects on neonate gut homeostasis cannot be excluded.
n2:mentions: n3:12603850

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rdf:value: antibacterial activity, cathelicidin-derived antimicrobial peptides have also been noted to exert other biological activities so as to promote cell differentiation and to stimulate angiogenesis and epithelial cell proliferation (15, >>16<<). Although our results do not support an involvement of epithelial CRAMP in the postnatal intestinal organ development, other CRAMP-mediated biological effects on neonate gut homeostasis cannot be excluded.
n2:mentions: n3:16177113

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rdf:value: Similar results have been found for cryptdin expression in Paneth cells (>>11<<). The close temporal association between the loss of CRAMP expression and developmental changes such as the increase of epithelial proliferation, the emergence of crypts, and the appearance of mature Paneth cells instead indicate a
n2:mentions: n3:11010975

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rdf:value: Indeed, cathelicidin expression has previously been linked to cell differentiation and exposure to differentiation-promoting agents (>>35<<, 36). Developmental regulation is also supported by the recently identified differential expression profile of the Wnt transcriptional effectors Tcf3 and Tcf4 during the development of the mouse intestinal epithelium (18). The identified
n2:mentions: n3:16497887

Subject Item: _:vb3044284
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rdf:value: Indeed, cathelicidin expression has previously been linked to cell differentiation and exposure to differentiation-promoting agents (35, >>36<<). Developmental regulation is also supported by the recently identified differential expression profile of the Wnt transcriptional effectors Tcf3 and Tcf4 during the development of the mouse intestinal epithelium (18). The identified
n2:mentions: n3:16740661

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rdf:value: Developmental regulation is also supported by the recently identified differential expression profile of the Wnt transcriptional effectors Tcf3 and Tcf4 during the development of the mouse intestinal epithelium (>>18<<). The identified increase in epithelial proliferation during the postnatal development is paralleled by enhanced Wnt signaling in the area of cellular proliferation (18). The restriction of CRAMP expression to neonate epithelium and the
n2:mentions: n3:17681174

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rdf:value: The identified increase in epithelial proliferation during the postnatal development is paralleled by enhanced Wnt signaling in the area of cellular proliferation (>>18<<). The restriction of CRAMP expression to neonate epithelium and the gradual loss of CRAMP-positive cells along the crypt–villus axis might therefore result from a combined developmental program of cell differentiation and proliferation.
n2:mentions: n3:17681174

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