Postnatal development and LPS responsiveness of pulmonary adenosine receptor expression and of adenosine-metabolizing enzymes in mice

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1 nture pulishing group Bsic Science Investigtion Postntl development nd responsiveness of pulmonry denosine receptor expression nd of denosine-metolizing enzymes in mice Juhni Metsol 1, Mikel Mksimow 2, Mrj Ojniemi 1, Hnn Metsol 3, Fumiko Mrttil-Ichihr 2, Reett Vuolteenho 1,4, Genndy G. Yegutkin 2, Mrko Slmi 2, Mikko Hllmn 1 nd Sirp Jlknen 2 Bckground: Adenosine levels re regulted y ecto- 5' -nucleotidse/cd73 nd denosine deminse (ADA). Adenosine regultes endothelil permeility nd ntiinflmmtory responses vi denosine receptors. Here, the denosine receptors nd purine-converting enzymes were studied during postntl development nd inflmmtion. Methods: Neworn, 1-, 10-, nd dult C57BL/6 mice were chllenged intrperitonelly (i.p.) with lipopolyscchride () for 6 h. The inflmmtory response ws evluted y histochemistry. Expression levels of denosine receptors (A1, A2A, A2B, nd A3), CD73, nd ADA were mesured y quntittive reverse trnscription polymerse chin rection. A1 ws studied y immunohistochemistry, nd enzyme ctivities were nlyzed y thin-lyer chromtogrphy. Results: cused respirtory distress in neworns within 24 h. induced neutrophils t the sl stge nd lveolr congestion. Low ctivity nd expression of CD73 incresed fter irth. Expression of ADA fter incresed 16-fold in dults nd 2-fold in neworns (P < 0.05). A1 expression ws high in neworns nd incresed fter (P < 0.05). A1 ws loclized to endothelil memrnes. A2A decresed fter in neworns nd incresed in dults (P < 0.05). The expression of A3 incresed in neworns nd dults fter. Conclusion: Low pulmonry CD73 expression, -induced suppression of A2A, -induced increse of A1 expression, nd severe respirtory distress were distinguishing responses in the neworns from those in the dults. Adenosine is purine se, which is n integrl component of DNA-RNA nd hs severl dditionl functions. It hs een shown to regulte vsculr endothelil integrity. As predominntly extrcellulr signling molecule, denosine hs recently een implicted s n importnt meditor of pulmonry inflmmtion (1). Ecto-5 nucleotidse (CD73, EC ) is cell-surfce enzyme, which produces extrcellulr denosine from denosine monophosphte (AMP) in the vsculture (2,3). Adenosine then inds to its receptors or is metolized y denosine deminse (ADA, EC ) or y denosine kinse (AK) to inosine or AMP, respectively (2). CD73 increses nd ADA decreses the denosine levels. It hs een shown tht interferon-β (IFN-β) enhnces the ctivity of CD73 in mice nd mn, leding to n increse in denosine levels in the vsculr endothelium nd decrese in vsculr lekge (4,5). The effects of denosine cn e nti- or proinflmmtory, depending on its concentrtion nd on the type of receptor eing ctivted. There re four different receptors tht trnsmit the effects of denosine to intrcellulr comprtments: A1, A2A, A2B, nd A3. A2AR nd A2BR re Gs-type G-proteincoupled receptors (GPCRs), which increse the levels of intrcellulr camp, wheres A1R nd A3R re Gi-type GPCRs. Activtion of Gi GPCR leds to decrese in intrcellulr camp levels (6). Lipopolyscchride (), n integrl prt of the outer wll of Grm-negtive cteri, is cple of lunching proinflmmtory response. Severl lines of evidence suggest role for denosine receptors in modulting -induced pulmonry inflmmtion (7,8). Activtion of A3R hs een shown to reduce toxicity in murine model of lung inflmmtion y downregulting -induced influx of polymorphonucler cells to the lungs (9). In porcine monocytes, n denosine nlog, N-ethylcroxymidodenosine, downregultes tumor necrosis fctor (TNF)-α production nd upregultes interleukin (IL)-8 production fter stimultion, the effect eing medited minly y A2AR (10). hs lso een implicted in the induction of A2AR expression in murine mcrophges, this eing primrily medited y nucler fctor-κb (11). Preclinicl studies hve shown tht n A2AR gonist improves lung function fter cute lung dmge nd tht the timely ctivtion of A2AR limits the inflmmtion (12). There is growing evidence of the importnce of the denosine system in erly postntl life. Neontl lood CD73 ctivity ws reported to e higher thn ADA ctivity fvoring elevted denosine levels (13). The ADA ctivity ws shown to increse The second nd the third uthor nd the lst two uthors contriuted eqully to this work. 1 Deprtment of Peditrics, Institute of Clinicl Medicine nd Medicl Reserch Center, Oulu University Hospitl nd University of Oulu, Oulu, Finlnd; 2 MediCity Reserch Lortory, University of Turku, Turku, Finlnd; 3 Deprtment of Pthology, Oulu University Hospitl nd University of Oulu, Oulu, Finlnd; 4 Trnsgenic Core Fcility, Biocenter Oulu, University of Oulu, Oulu, Finlnd. Correspondence: Juhni Metsol (juhni.metsol@medsol.fi) Received 15 Jnury 2014; ccepted 30 June 2014; dvnce online puliction 1 Octoer doi: /pr Volume 76 Numer 6 Decemer 2014 Peditric Reserch 515

2 Metsol et l. during postntl development. Serum ADA levels in preterm infnts ffected y ronchopulmonry dysplsi were elevted in the erly postntl phse (14). The high levels of denosine in the lood of neontes inhiited cute TNF-α response (15). Cffeine hs een proposed to inhiit TNF-α production vi ntgonizing A1R (16). Cffeine tretment of preterm infnts with pne of premturity hs een shown to decrese the plsm levels of TNF-α nd IL-6 (17). Respirtory distress shortly fter irth nd in erly infncy is mjor cuse of chronic moridity in premture neontes. The inflmmtory insult is either primry cuse of lung disese or principl fctor influencing its progression (18,19). At term irth nd therefter, the irwys re reltively resistnt to endotoxins. In contrst, preterm infnts my e susceptile to the consequences of -induced inflmmtory injury (20,21). In ddition, peritonel infection nd inflmmtion due to Grmnegtive cteri is ssocited with serious crdiorespirtory distress fter preterm irth (22,23). Endothelil function plys crucil role in mny inflmmtory diseses in which vsculr permeility is one of the min pthogenic fctors (24 26). The mechnisms protecting ginst infections nd generlized inflmmtion my lso require defense functions tht re confined to the vsculr endothelium. Neworns re susceptile to respirtory distress induced y sepsis often cused y Grm-negtive cteri from the intestinl trct. Since denosine is known to e cple of modulting the pulmonry inflmmtion, we hypothesized tht the fctors supporting high denosine levels during perintl period would protect the mice from pulmonry inflmmtion nd respirtory distress. Therefore, we studied the expression of denosine producing, sensing, nd metolizing proteins in response to intrperitonel (i.p.) in different ge groups of mice. Here, we show evidence tht oth sl expression levels nd responsiveness of denosine-metolizing enzymes nd denosine receptors to re ge-dependent. We propose tht the immturity of the denosine signling system needs to e considered s fctor predisposing to vsculr inflmmtory injury in erly life. RESULTS Sensitivity to Endotoxin-Induced Respirtory Distress nd Pulmonry Dmge In i.p. dose response experiments, neworn mice were more susceptile to toxicity when compred with older nimls. The dose selected (0.5 µg, serotype 0111:B4/g ody wt.) cused no detectle symptoms in the dult nimls (unpulished dt) nd resulted in no mortlity nd hrdly ny respirtory distress in pups (n = 7). In contrst, 53% of the neworns (n = 15; ge <12 h when ws given) presented with gsping nd skin color chnges nd died, indicting severe respirtory distress within h fter the insult. The rest of the neworn pups presented with respirtory distress tht ws not ftl during the oservtion period of 24 h. None of the plceo-injected pups (n = 8) developed respirtory distress. As the nimls ecme symptomtic within the first h fter the injection, in the susequent experiments, we scrificed the mice 6 h fter the injection to investigte the erly events. Histologicl nlyses of the lungs of neworn, nd dult mice were crried out to study the effects of. As expected, the structurlly immture lungs of neworn mice reveled no true lveoli, wheres lungs from nimls reveled lveolr sccules. The study hd power to detect only lrge differences in the inflmmtory response. However, only few trends in lung histology were evident, nd they did not rech sttisticl significnce (Tle 1): Neworn mice tended to hve more neutrophils t the sl stge thn the older nimls. Trnsmigrtion of neutrophils fter the endotoxin chllenge tended to increse neutrophil counts in 14-dold nd dult nimls, nd lveolr congestion incresed fter the insult especilly in the younger ge groups. The Ontogenic Chnges in CD73 nd ADA Activities There were no differences etween tretment groups in CD73 ctivity levels, nd therefore, we pooled control nd nimls to study the ontogenic differences. The ctivity of CD73 in the lungs incresed significntly with ge (Figure 1). In 10-d-old mice, CD73 ctivity ws threefold higher thn in the neworn mice (P < 0.05). To study whether the expression levels of CD73 prllel the chnges in its ctivity, CD73 mrna levels were mesured in neworn, nd dult mice. The expression levels of CD73 in the lungs were five- to sixfold higher in nd dult mice thn in neworn mice (P < 0.05) (Figure 2). The expression of CD73 did not differ etween nd dult mice. Since ADA ffects the serum levels of denosine, its expression levels were mesured s well. ADA ctivity in 10-d-old mice s compred to younger nimls ws twofold (Figure 1). Bsl expression levels of ADA reveled no detectle trends fter irth, either (Figure 2). The Ontogenic Chnges in Adenosine Receptor Expression Levels The expression levels of A1R, A2AR, A2BR, nd A3R in the lungs were mesured in neworn, nd dult mice. The expression levels of A1R re shown in Figure 3. The sl level ws twofold higher in neworn thn in nd dult mice (P < 0.05). The pulmonry expression levels of A2AR, A2BR, nd A3R did not show differences etween ge groups (Figure 3 d). Tle 1. Histologicl nlysis of lungs Neutrophil count per 5 high-power fields (SD) Alveolr congestion (SD) Age Shm Shm Neworn (34.9) (23.6) 1.7 (0.8) 2.4 (1.1) 78.3 (30.3) (45.1) 1.7 (0.8) 2.2 (0.8) 62 (30.4) (57.6) 1.3 (0.8) 1.7 (0.6) Alveolr congestion ws grded vi four-tiered system from two tissue sections: no congestion = 0, congestion in <25% of the field re = 1, in 25 49% of the field re = 2, in 50 74% of the field re = 3, nd in % of the field re = 4., lipopolyscchride. 516 Peditric Reserch Volume 76 Numer 6 Decemer 2014

3 Postntl denosine receptor responses CD73 ctivity (nmol/mg/hour) Reltive expression of CD ADA ctivity (nmol/mg/hour) Neworn 1 d old 10 d old Neworn 1 d old 10 d old Reltive expression of denosine deminse Neworn Neworn Figure 1. Enzyme ctivities of CD73 nd denosine deminse (ADA) increse with ge. Activity of CD73 () nd ADA () in 0-d-old neworn (n = 6), 1-d- (n = 6), nd 10-d-old (n = 4) nimls. P < The Expression Levels nd Activities of CD73 nd ADA After When the influence of i.p. chllenge on the expression levels of denosine-metolizing enzymes ws studied, did not lter the expression levels of CD73 in ny ge group (Figure 2). In contrst, the expression of ADA incresed 2- (neworn, ) to 16-fold (dult) fter the chllenge (P < 0.05). After the, the levels were eightfold higher in dult mice thn in neworn nd mice (P < 0.05). There were no detectle increses in either CD73 or ADA ctivities s studied 6 h fter the i.p. injection (dt not shown). The Expression Levels of Adenosine Receptors After There ws twofold increse in the expression of A1R fter injection in neworn nd mice compred with sline-injected nimls (P < 0.05). mice showed no increse of A1R expression fter the chllenge (Figure 3). In neworn mice, the expression of A2AR decresed 50% (P < 0.05) fter the injection, wheres in nimls, there were no detectle chnges in the expression levels fter. In dult mice, sixfold increse in A2A receptor expression ws detected fter the chllenge (P < 0.05). Overll, Neworn Neworn Figure 2. Expression levels of pulmonry CD73 nd denosine deminse (ADA) in nd lipopolyscchride () stimulted neworn,, nd dult mice. Reltive mrna expression levels of the enzymes CD73 () nd ADA () in neworn (n = 9 nd n = 10 in nd groups, respectively), (n = 6 nd n = 8 in nd groups, respectively) nd dult nimls (n = 5 in oth nd groups). P < the expression levels fter the insult were lower in neworn mice when compred with nd dult mice (P < 0.05). There were no detectle differences in -induced expression levels of A2BR etween the three ge groups (Figure 3,c). In neworn nd dult mice, the expression levels of A3R incresed fourfold fter the injection (P < 0.05). In nimls, the levels tended to e higher in the group. Expression levels of A3R fter the injection tended to e higher in neworn thn in mice (P < 0.05) (Figure 3d). The Immunohistochemistry of A1R Immunohistochemistry ws performed to study the distriution nd content of the A1R protein. A1R positivity ws evident in lungs from mny neworn (four out of seven nimls) (Figure 4) nd some nimls (two out of six) Volume 76 Numer 6 Decemer 2014 Peditric Reserch 517

4 Metsol et l. c d Figure 3. Expression levels of denosine receptors in nd lipopolyscchride () stimulted neworn,, nd dult mice. Reltive mrna expression levels of denosine receptors A1, A2A, A2B, nd A3 ( d, respectively) in neworn (n = 9 nd n = 10 in nd groups, respectively), (n = 6 nd n = 8 in nd groups, respectively), nd dult nimls (n = 5 in oth nd groups). P < (Figure 4d) 6 h fter the injection, wheres immunostining ws rely detectle in the lungs from neworn (two out of six) nd (two out of six) nimls (Figure 4,c). The stining ws detected minly in the endothelil cells. DISCUSSION In the present study, we found evidence of the vulnerility of structurlly immture lung tissue to vsculr inflmmtory insult, suggesting tht resistnce to i.p. develops s function of the postntl ge. In n ttempt to clrify the ge-dependent increse in tolernce to n inflmmtory insult, we studied the expression of oth the denosine receptors nd the expression nd ctivities of enzymes involved in denosine metolism, known to ply role in pulmonry immunity nd the mintennce of vsculr integrity (24,27,28). We found tht CD73 ctivity, protective ginst vsculr inflmmtion, incresed mrkedly soon fter irth. In ddition, the -medited induction of inflmmtory A1R expression decresed fter irth, wheres -induced suppression of nti-inflmmtory A2AR ws olished fter irth. We propose tht extrcellulr denosine levels nd denosine receptors my ply role in -induced toxicity. In the present study, we demonstrte neontl increse in oth the ctivity nd the expression of CD73, suggesting n increse in the cpcity to generte extrcellulr denosine. It my e rgued tht neworn mice hve lower cpcity to produce denosine, resulting in stronger -induced inflmmtory responses compred to older nimls. In contrst, the expression of Toll-like receptor 4 (29,30) nd the inflmmtory cytokine responses (31) increse during the perintl development, ut do not prllel the perintl development of resistnce ginst cute inflmmtory insults. The sl expression levels of proinflmmtory A1R were higher in neworn mice nd the -induced increse in the expression levels ws evident only in the neworn mice. A1R protein ws detected in endothelil cell memrnes prticulrly fter dministrtion of. The differences seen in A1R expression etween neworn nd older mice my e one 518 Peditric Reserch Volume 76 Numer 6 Decemer 2014

5 Postntl denosine receptor responses c d Figure 4. increses denosine receptor A1 immunostining compred to mice. Immunohistochemistry stining using A1R ntiody with hemtoxylin-eosin counterstining of neworn lungs (), neworn lungs 6 h fter the lipopolyscchride () exposure (), nd lungs (c) nd mouse lungs 6 h fter the exposure (d). Bottom right r = 100 μm. mechnism explining the inflmmtory stte in the neworns. The sl expression levels of denosine receptors A2AR, A2BR, nd A3R reveled no remrkle trends in postntl lung. However, the expression levels in response to were different etween the ge groups (Tle 2). Most strikingly, the nti-inflmmtory A2AR reveled -induced suppression in neworns, wheres in dults, significnt -induced increse of A2AR expression ws evident. The immunohistochemicl verifiction of these findings filed due to low qulity of ville commercil A2AR ntiodies. Adenosine deminse hs een considered to e n llosteric gonist for A1R, nd A2AR (32) nd ADA / mice hve chronic pulmonry inflmmtion nd firosis (33). This rises the possiility tht interction etween ADA nd A2AR my influence the inflmmtory response. However, direct evidence for this is lcking. In the present study, we found no detectle increse in ADA ctivity within 6 h fter i.p.. However, rpidly induced n increse in the expression of ADA in ech ge group. In dult mice, the increse ws ~16-fold, compred with 2-fold increse in neworn mice (Figure 2). The discrepncy etween the mrna levels nd enzymtic ctivity of ADA my e explined y the dely etween the mrna induction nd the protein synthesis. An increse in ADA without n increse in CD73 my hve n dditionl influence on the dverse consequences of in the neworn. The intriguing possiility tht serum denosine levels influence the resistnce to remins to e studied. Acute lung injury induced y is well documented nd previous results re consistent with the present oservtions (34,35). Within 24 h fter i.p. dministrtion of, most neworn pups developed signs of severe respirtory distress. In contrst, the sme dose given to nimls did not cuse evidence Tle 2. Chnges in gene expression fter injection Neworn Shm Shm Shm CD73 Low N.C. High N.C. High N.C. ADA A1R High Low Low N.C. A2AR N.C. A2BR N.C. N.C. N.C. A3R N.C. = injection incresed expression levels. = The increse ws significntly greter thn other increse(s) in sme direction. = injection decresed expression levels., lipopolyscchride; N.C., no chnges. of respirtory distress. Study of lung histology 6 h fter injection reveled trend towrd more severe lung injury in neworn compred with older mice. However, the neutrophil count in the neworn mice fter chllenge ws comprle to tht in the older mice. In study y Alvir et l. (36), the inflmmtory response fter i.p. ws milder nd the dverse consequences to ctivtion of nucler fctor-κb were fewer in 5-d-old mice thn in dults. We propose tht the increse in resistnce of the vsculr endothelium ginst inflmmtory injury my depend on the development of responsiveness of the denosine receptors nd enzymtic ctivities involved in denosine metolism. In conclusion, our results show evidence tht compred to dult mice, neworn mice hve significntly lower expression of pulmonry nti-inflmmtory denosine receptors nd significntly incresed expression of proinflmmtory denosine receptors fter n inflmmtory insult. In ddition, the expression of ADA during the inflmmtory stte ws lower in the Volume 76 Numer 6 Decemer 2014 Peditric Reserch 519

6 Metsol et l. Tle 3. Primer sequences for quntittive polymerse chin rection Gene Protein Sequence 5-3 Finl concentrtion (nmol/l) Product size (p) Ador1 A1R F: CAA GGG AGA GAA TCC AGC AG; R: ACC TCC GAG TCA AGA TCC CT Ador2 A2AR F: CCT TCA TAC CCG TCA CCA AG; R: GCA GAG TTC CAT CTT CAG CC Ador2 A2BR F: TGT CCC AGT GAC CAA ACC TT; R: TGC TCA CAC AGA GCT CCA TC Ador3 A3R F: GGA AAC TAG CCA GCA AAG GC; R: TGC TGT ACA CCG ATA CCT GC ADA ADA F: GAT GGT TTC TGG CTT GAT GG; R: ACG CAG ACC CAG AGA GCT T NT5E CD73 F: GTC CCC CAT TGA TGA GAA GA; R: CAA AAG CCT TCT TCA GGG TG TBP TBP F: GAA TAT AAT CCC AAG CGA TTT G; R: CAC ACC ATT TTT CCA GAA CTG TBP, TATA-ox inding protein. neworn nd young nimls thn in dults. ADA hs een shown to increse the ffinity of the A2A receptor to denosine (37). Finlly, CD73, ctlyzing the formtion of denosine, incresed in prllel with vsculr development of murine lung shortly fter irth. In humn, lung nlogous vsculr development strts preterm (38). We propose tht the incresing resistnce to vsculr inflmmtory insults my e influenced y denosine receptors nd enzymes regulting oth denosine levels nd its ffinity to specific receptors. The immture inflmmtory responses of ADA nd the denosine receptors in preterm infnts susceptile to chronic lung disese remin to e studied. METHODS Animls Neworn (ge <24 h t the onset of the study), 1-d-old, 10-d-old,, nd dult C57BL/6 mice from the Lortory Animl Center of University of Oulu were used. All studies were pproved y the Finnish Animl Ethics Committee. Mouse Model of i.p. Escherichi coli (serotype 0111:B4, Sigm-Aldrich, St Louis, MO) or 0.9% sline crrier ws injected i.p. s single olus using NovoPen junior insulin injector with NovoFine 32G needle (NovoNordisk A/S, Bgsverd, Denmrk). In preliminry experiments, the mice were inspected every 2 3 h during the experiment for 24 h. In cses of severe respirtory distress (cynosis, gsping), the mice were euthnized. The dose chosen provoked respirtory filure in 50 percent of the nimls in pilot studies using neworn mice (dt not shown). The dose ws 0.5 µg/g for ech niml. The concentrtion of reconstituted ws 0.03 µg/µl. The specimens were collected 6 h fter the i.p. or sline. Before collection, the nimls were euthnized y cutting the neck verter. The chest ws opened y medin sternotomy. Cre ws tken to void leeding to the trche during lung preprtion. The lungs were immeditely frozen in liquid nitrogen nd stored t 74 C or fixed in 4% formldehyde. Anlysis of Expression of CD73, ADA, nd Adenosine Receptors Totl RNA from lungs ws isolted using Tri-regent (Sigm-Aldrich) ccording to the mnufcturer s instructions. For quntittive reverse trnscription polymerse chin rection, the RNA ws purified y using RNesy Micro Kits (Qigen Venlo, NL). cdna ws cquired from the purified mrna with iscript cdna Synthesis Kit (Bio-Rd Lortories, Hercules, CA). Levels of gene expression of CD73 (NT5E), ADA, nd denosine receptors A1 (Ador1), A2A (Ador2), A2B (Ador 2), nd A3 (Ador3) were mesured y using Bio-Rd 520 Peditric Reserch Volume 76 Numer 6 Decemer p 250 p Lne A B C D E F G Figure 5. Gel electrophoresis of the polymerse chin rection products. TBP, CD73, ADA, A1R, A2AR, A2BR, nd A3R re in lnes g, respectively. 250 p stndrd ws used. Cfx-96 rel-time PCR detection system (Bio-Rd Lortories). Reltive quntifiction ws crried out with the ΔC t method using the TATA-inding protein gene s reference gene. The primers used re shown in Tle 3. The sizes of PCR products were confirmed y mens of gel electrophoresis in which the 2% grose in 1 TAE ws used (Figure 5). The numer of independent nlyses ws four to six for ech individul group of specimens. Anlysis of the CD73 nd ADA Activities CD73 nd ADA ctivities were mesured in lung tissue lystes from the neworn, 1-d-old nd 10-d-old mice using thin lyer chromtogrphy method previously descried y Yegutkin et l. (39). Briefly, tissue lystes (20 μg of protein) were incuted for 60 min t 37 C in finl volume of 80 µl RPMI-1640 medium contining 4 mmol/l β-glycerophosphte nd 300 μmol/l of either [2-3 H]AMP (Quotient Bioreserch, GE Helthcre, Rushden, UK) or [2-3 H]denosine (Amer, Little Chlfont, UK) in the cse of ecto-5 -nucleotidse/cd73 nd ADA ssys, respectively. Ctlytic rections were terminted y pplying liquots of the mixture onto Alugrm SIL G/UV 254 sheets (Mcherey-Ngel, Duren, Germny). 3 H-leled AMP nd nucleosides were seprted y TLC nd quntified y scintilltion β-counting. Histology nd Immunohistohemistry Tissue smples were emedded in prffin nd cut into 2.5-µm sections. The sections were stined with hemtoxylin-eosin. Neutrophil counts were ssessed in five high-power fields. In ddition, lveolr congestion ws grded on four-tiered systems from two tissue sections: no congestion = 0, congestion in <25% of the field re = 1, in 25 49% of the field re = 2, in 50 74% of the field re = 3, nd % of the field re = 4 (40). The histologicl exminer ws linded s to which group the niml elonged to. For the immunohistochemicl stining of A1R in mouse lungs, 3.5 µm sections were tken from formlin-fixed prffin-emedded tissue nd the rit polyclonl nti-adora1 ntiody from Acm (82477, Acm, Cmridge, MA) ws used.

7 Postntl denosine receptor responses Sttisticl Anlysis One-wy ANOVA with two-smple t-tests s post hoc tests were used for comprison of mens. Bonferroni corrections were used for multiple comprisons. The level of significnce ws P < OriginPro v softwre (OriginL, Northmpton, MA) ws used for sttisticl nlysis. ACKNOWLEDGMENTS We thnk Mrit Hrl, Riitt Vuento, Sri Mäki, nd Erj Tomperi for technicl ssistnce. STATEMENT OF FINANCIAL SUPPORT This study ws supported y the Alm nd K. A. Snellmn Foundtion, Oulu, Finlnd; the Foundtion for Peditric Reserch, Helsinki, Finlnd; Sigrid Juselius Foundtion, Helsinki, Finlnd; nd the Acdemy of Finlnd, Helsinki, Finlnd. Disclosure: The uthors hve no conflict of interest to disclose. References 1. Schepp CP, Reutershn J. Bench-to-edside review: denosine receptors promising trgets in cute lung injury? Crit Cre 2008;12: Yegutkin GG. Nucleotide- nd nucleoside-converting ectoenzymes: Importnt modultors of purinergic signlling cscde. Biochim Biophys Act 2008;1783: Zimmermnn H, Zeisch M, Sträter N. Cellulr function nd moleculr structure of ecto-nucleotidses. Purinergic Signl 2012;8: Kiss J, Yegutkin GG, Koskinen K, Svunen T, Jlknen S, Slmi M. IFNet protects from vsculr lekge vi up-regultion of CD73. Eur J Immunol 2007;37: Bellingn G, Mksimow M, Howell DC, et l. The effect of intrvenous interferon-et-1 (FP-1201) on lung CD73 expression nd on cute respirtory distress syndrome mortlity: n open-lel study. Lncet Respir Med 2014;2: Krmouty-Quintn H, Xi Y, Blckurn MR. Adenosine signling during cute nd chronic disese sttes. J Mol Med (Berl) 2013;91: He X, Hu JL, Li J, et l. A feedck loop in PPARγ-denosine A2A receptor signling inhiits inflmmtion nd ttenutes lung dmges in mouse model of -induced cute lung injury. Cell Signl 2013;25: Li J, Zho L, He X, Zeng YJ, Di SS. Sinomenine protects ginst lipopolyscchride-induced cute lung injury in mice vi denosine A(2A) receptor signling. PLoS One 2013;8:e Wgner R, Ngmsri KC, Strk S, Vollmer I, Reutershn J. Adenosine receptor A3 is criticl meditor in -induced pulmonry inflmmtion. Am J Physiol Lung Cell Mol Physiol 2010;299:L Ondrckov P, Kovru H, Kovru F, Lev L, Fldyn M. Adenosine modultes -induced cytokine production in porcine monocytes. Cytokine 2013;61: Elson G, Eisenerg M, Grg C, et l. Induction of murine denosine A(2A) receptor expression y : nlysis of the 5 upstrem promoter. Genes Immun 2013;14: Folkesson HG, Kuzenko SR, Lipson DA, Mtthy MA, Simmons MA. The denosine 2A receptor gonist GW328267C improves lung function fter cute lung injury in rts. Am J Physiol Lung Cell Mol Physiol 2012;303:L Pettengill M, Roson S, Tresenriter M, et l. Solule ecto-5 -nucleotidse (5 -NT), lkline phosphtse, nd denosine deminse (ADA1) ctivities in neontl lood fvor elevted extrcellulr denosine. J Biol Chem 2013;288: Cnpolt FE, Yurdkök M, Korkmz A, Yiğit S, Tekinlp G. Adenosine deminse levels in premture infnts with respirtory distress syndrome nd ronchopulmonry dysplsi. J Mtern Fetl Neontl Med 2011;24: Levy O, Coughlin M, Cronstein BN, Roy RM, Desi A, Wessels MR. The denosine system selectively inhiits TLR-medited TNF-lph production in the humn neworn. J Immunol 2006;177: Chvez-Vldez R, Wills-Krp M, Ahlwt R, Cristoflo EA, Nthn A, Gud EB. Cffeine modultes TNF-lph production y cord lood monocytes: the role of denosine receptors. Peditr Res 2009;65: Chvez Vldez R, Ahlwt R, Wills-Krp M, Nthn A, Ezell T, Gud EB. Correltion etween serum cffeine levels nd chnges in cytokine profile in cohort of preterm infnts. J Peditr 2011;158:57 64, 64.e Joe AH. Effects of choriomnionitis on the fetl lung. Clin Perintol 2012;39: Speer CP. Choriomnionitis, postntl fctors nd proinflmmtory response in the pthogenetic sequence of ronchopulmonry dysplsi. Neontology 2009;95: Yerkovich ST, Wikström ME, Suriyrchchi D, Prescott SL, Uphm JW, Holt PG. Postntl development of monocyte cytokine responses to cteril lipopolyscchride. Peditr Res 2007;62: Kunzmnn S, Collins JJ, Kuypers E, Krmer BW. Thrown off lnce: the effect of ntentl inflmmtion on the developing lung nd immune system. Am J Ostet Gynecol 2013;208: Adms-Chpmn I. Long-term impct of infection on the preterm neonte. Semin Perintol 2012;36: Reiss LK, Uhlig U, Uhlig S. Models nd mechnisms of cute lung injury cused y direct insults. Eur J Cell Biol 2012;91: Ålgrs A, Krikoski M, Yegutkin GG, et l. Different role of CD73 in leukocyte trfficking vi lood nd lymph vessels. Blood 2011;117: Stevens T. Functionl nd moleculr heterogeneity of pulmonry endothelil cells. Proc Am Thorc Soc 2011;8: Zhng Q, Coveney AP, Yu S, et l. Inefficient ntimicroil functions of innte phgocytes render infnt mice more susceptile to cteril infection. Eur J Immunol 2013;43: Petrovic-Djergovic D, Hymn MC, Ry JJ, et l. Tissue-resident ecto-5 nucleotidse (CD73) regultes leukocyte trfficking in the ischemic rin. 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