diff --git a/grobid-trainer/resources/dataset/segmentation/corpus/raw/pub.1158465915.training.segmentation b/grobid-trainer/resources/dataset/segmentation/corpus/raw/pub.1158465915.training.segmentation new file mode 100644 index 0000000000..8ce34d2bbc --- /dev/null +++ b/grobid-trainer/resources/dataset/segmentation/corpus/raw/pub.1158465915.training.segmentation @@ -0,0 +1,931 @@ +Title Title title T Ti Tit Titl BLOCKSTART PAGESTART NEWFONT HIGHERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 0 no 0 10 0 1 0 0 0 +Patient-specific alterations patient-specific P Pa Pat Pati BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 0 - 1 10 0 0 0 0 1 +cancer patients cancer c ca can canc BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 0 no 0 10 0 0 0 0 1 +Authors Authors authors A Au Aut Auth BLOCKSTART PAGEIN NEWFONT HIGHERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 0 no 0 10 0 0 0 0 0 +Annelien Morlion annelien A An Ann Anne BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 0 1 (,),(,,),(,,), 14 10 0 0 0 0 1 +Anckaert (1,2), anckaert A An Anc Anck BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 0 2 (,),(,),(,),(,), 16 10 0 0 0 0 1 +Celine Everaert celine C Ce Cel Celi BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 1 0 0 0 0 0 0 0 3 (),(),(),(,), 13 10 0 0 0 0 0 +Mestdagh°* (1,2) mestdagh°* M Me Mes Mest BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 0 4 *(,) 4 10 0 0 0 0 0 +(1) Department (1) ( (1 (1) (1) BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 0 4 (),,, 5 10 0 0 0 0 0 +(2) OncoRNALab, (2) ( (2 (2) (2) BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 0 5 (),(),, 7 10 0 0 0 0 0 +(3) Department (3) ( (3 (3) (3) BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 0 5 (),,, 5 10 0 0 0 0 0 +(4) TOBI (4) ( (4 (4) (4) BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 0 6 (),,-,, 7 10 0 0 0 0 0 +(5) Department (5) ( (5 (5) (5) BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 0 7 (),,, 5 10 0 0 0 0 0 +°Both authors °both ° °B °Bo °Bot BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 0 7 no 0 10 0 0 0 0 0 +*Corresponding author *corresponding * *C *Co *Cor BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 0 8 * 1 10 0 0 0 0 0 +Author contributions author A Au Aut Auth BLOCKSTART PAGEIN NEWFONT HIGHERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 8 no 0 10 0 0 0 0 0 +Conceptualization: AM, conceptualization: C Co Con Conc BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 0 8 :,,, 4 10 0 0 0 0 0 +Data Curation: data D Da Dat Data BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 8 : 1 10 0 0 0 0 0 +Funding acquisition: funding F Fu Fun Fund BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 9 :,,, 4 10 0 0 0 0 0 +Software: AM, software: S So Sof Soft BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 9 :, 2 10 0 0 0 0 0 +Validation: AM validation: V Va Val Vali BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 9 : 1 10 0 0 0 0 0 +Formal analysis: formal F Fo For Form BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 9 : 1 10 0 0 0 0 0 +Investigation: EV, investigation: I In Inv Inve BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 10 :,, 3 10 0 0 0 0 0 +Methodology: AM, methodology: M Me Met Meth BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 10 :,, 3 10 0 0 0 0 0 +Project administration: project P Pr Pro Proj BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 10 :,, 3 10 0 0 0 0 0 +Resources: FO, resources: R Re Res Reso BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 11 :,,,,, 6 10 0 0 0 0 0 +Supervision: JV, supervision: S Su Sup Supe BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 11 :, 2 10 0 0 0 0 0 +Visualization: AM visualization: V Vi Vis Visu BLOCKSTART PAGEEND SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 0 11 : 1 10 0 0 0 0 0 +Writing -original writing W Wr Wri Writ BLOCKSTART PAGESTART SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 0 -:,, 4 10 0 1 0 0 1 +Writing -review writing W Wr Wri Writ BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 0 -: 2 10 0 1 0 0 1 +Abstract Abstract abstract A Ab Abs Abst BLOCKSTART PAGEIN NEWFONT HIGHERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 0 no 0 10 0 0 0 0 0 +Circulating nucleic circulating C Ci Cir Circ BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 0 no 0 10 0 0 0 0 1 +and disease. and a an and and BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 0 .,- 3 10 0 0 0 0 1 +266 cancer 266 2 26 266 266 BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS ALLDIGIT 0 0 0 0 0 0 0 0 0 1 -(,; 4 10 0 0 0 0 1 +n=58, 3 n=58, n n= n=5 n=58 BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 0 1 ,).--- 6 10 0 0 0 0 1 +transcriptomes compared transcriptomes t tr tra tran BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 0 2 . 1 10 0 0 0 0 0 +among patients among a am amo amon BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 2 ,. 2 10 0 0 0 0 0 +Therefore, we therefore, T Th The Ther BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 3 , 1 10 0 0 0 0 0 +reference control reference r re ref refe BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 3 -. 2 10 0 0 0 0 0 +genes discriminate genes g ge gen gene BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 0 4 ,, 2 10 0 0 0 0 0 +accuracy (AUC accuracy a ac acc accu BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 4 (.). 4 10 0 0 0 0 0 +(2 lymphoma (2 ( (2 (2 (2 BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 0 5 ()()., 6 10 0 0 0 0 0 +demonstrate heterogeneity demonstrate d de dem demo BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 5 - 1 10 0 0 0 0 0 +that case-specific that t th tha that BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 5 -. 2 10 0 0 0 0 0 +Introduction Introduction introduction I In Int Intr BLOCKSTART PAGEIN NEWFONT HIGHERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 6 no 0 10 0 0 0 0 0 +Cancer is cancer C Ca Can Canc BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 6 , 1 10 0 0 0 0 0 +cancer types. cancer c ca can canc BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 6 . 1 10 0 0 0 0 0 +well-being and well-being w we wel well BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 7 -., 3 10 0 0 0 0 0 +enabling treatment enabling e en ena enab BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 7 ,, 2 10 0 0 0 0 0 +selection or selection s se sel sele BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 8 . 1 10 0 0 0 0 0 +biopsies, liquid biopsies, b bi bio biop BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 8 ,- 2 10 0 0 0 0 0 +promising tool promising p pr pro prom BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 8 . 1 10 0 0 0 0 0 +To date, to T To To To BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 9 , 1 10 0 0 0 0 0 +circulating tumor circulating c ci cir circ BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 9 (). 3 10 0 0 0 0 0 +on the on o on on on BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 10 , 1 10 0 0 0 0 0 +cancer types cancer c ca can canc BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 10 -.-() 5 10 0 0 0 0 0 +reflecting dynamic reflecting r re ref refl BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 11 , 1 10 0 0 0 0 0 +or upon or o or or or BLOCKSTART PAGEEND SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 11 ,. 2 10 0 0 0 0 0 +cell types cell c ce cel cell BLOCKSTART PAGESTART SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 0 -, 2 10 0 0 0 0 1 +allows monitoring allows a al all allo BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 0 --,- 4 10 0 0 0 0 1 +and Alzheimer's and a an and and BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 0 '. 2 10 0 0 0 0 1 +demonstrated for demonstrated d de dem demo BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 1 -. 2 10 0 0 0 0 1 +shown potential shown s sh sho show BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 1 -.. 3 10 0 0 0 0 1 +specifically looked specifically s sp spe spec BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 2 ""() 4 10 0 0 0 0 1 +healthy controls healthy h he hea heal BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 2 / 1 10 0 0 0 0 1 +indications of indications i in ind indi BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 2 --.. 4 10 0 0 0 0 1 +microbe-derived plasma microbe-derived m mi mic micr BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 3 - 1 10 0 0 0 0 0 +(colorectum, stomach, (colorectum, ( (c (co (col BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 0 3 (,,,,) 6 10 0 0 0 0 0 +group-level differences. group-level g gr gro grou BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 4 -.-. 4 10 0 0 0 0 0 +for liver for f fo for for BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 4 -, 2 10 0 0 0 0 0 +a higher a a a a a BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 1 0 1 0 0 0 0 0 0 4 - 1 10 0 0 0 0 0 +level differential level l le lev leve BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 5 .,, 3 10 0 0 0 0 0 +major contributors major m ma maj majo BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 1 1 0 0 0 0 0 0 5 . 1 10 0 0 0 0 0 +Here, we here, H He Her Here BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 5 , 1 10 0 0 0 0 0 +different cohorts different d di dif diff BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 6 ().- 4 10 0 0 0 0 0 +plasma from plasma p pl pla plas BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 6 - 1 10 0 0 0 0 0 +donors to donors d do don dono BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 7 --. 3 10 0 0 0 0 0 +Results revealed results R Re Res Resu BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 1 1 0 0 0 0 0 0 7 -, 2 10 0 0 0 0 0 +questioning the questioning q qu que ques BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 7 -.,- 4 10 0 0 0 0 0 +alterations, reflected alterations, a al alt alte BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 8 ,- 2 10 0 0 0 0 0 +patient sample patient p pa pat pati BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 8 -, 2 10 0 0 0 0 0 +distinguish cancer distinguish d di dis dist BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 8 . 1 10 0 1 0 0 0 +Table 1: table T Ta Tab Tabl BLOCKSTART PAGEIN NEWFONT LOWERFONT 1 1 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 8 :.,(),, 7 10 0 1 0 0 0 +method and method m me met meth BLOCKSTART PAGEIN NEWFONT SAMEFONTSIZE 0 1 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 9 .(*)-, 6 10 0 1 0 0 0 +data of data d da dat data BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 9 (). 3 10 0 1 0 0 0 +cohort name cohort c co coh coho BLOCKSTART PAGEIN NEWFONT LOWERFONT 1 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 0 10 no 0 7 0 1 0 0 0 +cancer patients cancer c ca can canc BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 10 no 0 10 0 1 0 0 0 +(cancer types) (cancer ( (c (ca (can BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 0 10 () 2 9 0 1 0 0 0 +controls biofluid controls c co con cont BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 10 no 0 9 0 1 0 0 0 +library preparation library l li lib libr BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 10 no 0 10 0 1 0 0 0 +application application application a ap app appl BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 10 no 0 6 0 1 0 0 0 +pan-cancer pan-cancer pan-cancer p pa pan pan- BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 10 - 1 10 0 1 0 0 0 +cohort cohort cohort c co coh coho BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 0 10 no 0 6 0 1 0 0 0 +200 (25) 200 2 20 200 200 BLOCKSTART PAGEIN NEWFONT SAMEFONTSIZE 0 0 NOCAPS ALLDIGIT 0 0 0 0 0 0 0 0 0 10 () 2 3 0 1 0 0 0 +8 8 8 8 8 8 8 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS ALLDIGIT 1 0 0 0 0 0 0 0 0 10 no 0 0 0 1 0 0 0 +plasma plasma plasma p pl pla plas BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 10 no 0 2 0 1 0 0 0 +mRNA capture mrna m mR mRN mRNA BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 0 10 ( 1 10 0 1 0 0 0 +Prep with prep P Pr Pre Prep BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 0 10 ) 1 8 0 1 0 0 0 +discovery discovery discovery d di dis disc BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 11 no 0 10 0 1 0 0 0 +three-cancer three-cancer three-cancer t th thr thre BLOCKSTART PAGEIN NEWFONT SAMEFONTSIZE 1 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 11 - 1 10 0 1 0 0 0 +cohort cohort cohort c co coh coho BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 0 11 no 0 5 0 1 0 0 0 +36 (3) 36 3 36 36 36 BLOCKSTART PAGEIN NEWFONT SAMEFONTSIZE 0 0 NOCAPS ALLDIGIT 0 0 0 0 0 0 0 0 0 11 () 2 2 0 1 0 0 0 +30* 30* 30* 3 30 30* 30* BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 0 11 * 1 1 0 1 0 0 0 +plasma plasma plasma p pl pla plas BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 11 no 0 2 0 1 0 0 0 +mRNA capture mrna m mR mRN mRNA BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 0 11 ( 1 10 0 1 0 0 0 +Prep with prep P Pr Pre Prep BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 0 11 ) 1 8 0 1 0 0 0 +validation pan-cancer validation v va val vali BLOCKSTART PAGEIN 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However, cohort. c co coh coho BLOCKSTART PAGEEND SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 3 11 ., 2 10 0 0 0 0 0 +pan-cancer cohort pan-cancer p pa pan pan- BLOCKSTART PAGESTART SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 0 --, 3 10 0 1 0 0 1 +(Figure 4d, (figure ( (F (Fi (Fig BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 3 0 (,). 4 10 0 1 0 0 1 +Cell type cell C Ce Cel Cell BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 1 ,/, 3 10 0 0 0 0 1 +monocytes and monocytes m mo mon mono BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 3 1 --(, 4 10 0 0 0 0 1 +Supplemental Table supplemental S Su Sup Supp BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 2 ),. 3 10 0 0 0 0 1 +The higher the T Th The The BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 3 - 1 10 0 0 0 0 1 +cancer cohort cancer c ca can canc BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 4 (--.-, 6 10 0 0 0 0 1 +0.679), especially 0.679), 0 0. 0.6 0.67 BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 5 .),()., 7 10 0 0 0 0 1 +erythrocyte/erythroid progenitor erythrocyte/erythroid e er ery eryt BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 3 6 /,- 3 10 0 0 0 0 0 +fraction in fraction f fr fra frac BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 7 (--.- 5 10 0 0 0 0 0 +(effect size=0.556), (effect ( (e (ef (eff BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 3 8 (.),.-(.),.-(.),.-( 19 10 0 0 0 0 0 +size=0.261), respectively). size=0.261), s si siz size BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 9 .),). 5 10 0 0 0 0 0 +Taken together, taken T Ta Tak Take BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 3 9 ,- 2 10 0 0 0 0 0 +cancer and cancer c ca can canc BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 10 -, 2 10 0 0 0 0 0 +and cell and a an and and BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 11 no 0 10 0 0 0 0 0 +cancer patients. cancer c ca can canc BLOCKSTART PAGEEND SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 11 . 1 10 0 0 0 0 0 +Figure 4: figure F Fi Fig Figu BLOCKSTART PAGESTART NEWFONT LOWERFONT 1 1 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 0 :--,, 5 10 1 1 0 0 0 +overlap in overlap o ov ove over BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 1 1 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 1 ., 2 10 1 1 0 0 0 +shows distinct shows s sh sho show BLOCKSTART PAGEIN NEWFONT SAMEFONTSIZE 0 1 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 2 ()., 4 10 1 1 0 0 0 +0verlap between 0verlap 0 0v 0ve 0ver BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 3 -. 2 10 1 1 0 0 0 +proportional to proportional p pr pro prop BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 4 4 ., 2 10 1 1 0 0 0 +control for control c co con cont BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 4 5 --.':.-,., 10 10 0 0 0 0 0 +index=8.3% for index=8.3% i in ind inde BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 6 .();.-,., 9 10 0 0 0 0 0 +index=14.3% for index=14.3% i in ind inde BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 8 .().., 6 10 0 0 0 0 0 +enrichment analysis enrichment e en enr enri BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 4 9 .. 2 10 0 0 0 0 0 +with q with w wi wit with BLOCKSTART PAGEEND SAMEFONT SAMEFONTSIZE 0 1 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 4 10 .()., 5 10 0 0 0 0 0 +abundant cell abundant a ab abu abun BLOCKSTART PAGESTART SAMEFONT SAMEFONTSIZE 0 1 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 4 0 .: 2 10 0 1 0 0 1 +types. 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0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 5 7 (:[.-.])(). 11 10 0 0 0 0 0 +We then we W We We We BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 5 7 no 0 10 0 0 0 0 0 +of biomarker of o of of of BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 5 8 .---, 5 10 0 0 0 0 0 +obtained perfect obtained o ob obt obta BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 5 8 no 0 10 0 0 0 0 0 +(AUC=1.000, based (auc=1.000, ( (A (AU (AUC BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 5 9 (.,,.), 7 10 0 0 0 0 0 +performance for performance p pe per perf BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 5 9 (., 3 10 0 0 0 0 0 +genes) (Figure genes) g ge gen gene BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 5 10 )(). 4 10 0 0 0 0 0 +samples (AUC=0.980, samples s sa sam samp BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 1 1 0 0 0 0 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The regulation. R Re Reg Regu BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 10 4 .-- 3 10 0 0 0 0 0 +EGAS00001006755 (dataset egas00001006755 E EG EGA EGAS BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 1 0 0 0 10 5 (). 3 10 0 1 0 0 0 +under study under u un und unde BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 10 5 (). 3 10 0 1 0 0 0 +FASTQs can fastqs F FA FAS FAST BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 10 6 (). 3 10 0 1 0 0 0 +Differential abundance differential D Di Dif Diff BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 10 6 () 2 10 0 1 0 0 0 +in a in i in in in BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 1 1 0 0 0 0 0 10 7 (:./.). 7 10 0 1 0 0 0 +Code availability code C Co Cod Code BLOCKSTART PAGEIN NEWFONT HIGHERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 10 7 no 0 10 0 1 0 0 0 +Code for code C Co Cod Code BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 10 7 no 0 10 0 1 0 0 0 +https://github.com/OncoRNALab/tailgenes. https://github.com/OncoRNALab/tailgenes. https://github.com/oncornalab/tailgenes. h ht htt http BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 1 10 7 ://.//. 7 10 0 1 0 0 0 +Acknowledgements Acknowledgements acknowledgements A Ac Ack Ackn BLOCKSTART PAGEIN NEWFONT HIGHERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 10 8 no 0 10 0 1 0 0 0 +Figures 1a figures F Fi Fig Figu BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 10 8 .. 2 10 0 1 0 0 0 +providing the providing p pr pro prov BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 10 8 , 1 10 0 1 0 0 0 +the first the t th the the BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 10 9 ,. 2 10 0 1 0 0 0 +Funding Funding funding F Fu Fun Fund BLOCKSTART PAGEIN NEWFONT HIGHERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 10 10 no 0 10 0 1 0 0 0 +This work this T Th Thi This BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 10 10 (;//), 6 10 0 1 0 0 0 +Hospital, Fund hospital, H Ho Hos Hosp BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 10 10 ,(;..,.., 9 10 0 0 0 0 0 +G0B2820N), and g0b2820n), G G0 G0B G0B2 BLOCKSTART PAGEEND SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 1 0 0 0 10 11 ),(,). 6 10 0 0 0 0 0 +Conflict of conflict C Co Con Conf BLOCKSTART PAGESTART NEWFONT HIGHERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 10 0 no 0 10 0 1 0 0 1 +A.M., J.V. a.m., A A. A.M A.M. BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 10 0 ..,..... 8 10 0 0 0 0 1 +References References references R Re Ref Refe BLOCKSTART PAGEIN NEWFONT HIGHERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 10 0 no 0 10 0 0 0 0 0 +1. Pessoa, 1. 1 1. 1. 1. BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 10 1 .,..,,.,..:. 12 10 0 0 0 0 1 +Critical Reviews critical C Cr Cri Crit BLOCKSTART PAGEIN NEWFONT SAMEFONTSIZE 0 1 INITCAP NODIGIT 0 0 1 0 0 0 0 0 10 1 /,(). 5 10 0 0 0 0 1 +2. Vorperian, 2. 2 2. 2. 2. BLOCKSTART PAGEIN NEWFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 10 2 .,..,,..,..- 12 10 0 0 0 0 1 +transcriptome. Nat transcriptome. t tr tra tran BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 10 3 .,-(). 6 10 0 0 0 0 0 +3. Moufarrej, 3. 3 3. 3. 3. BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 10 3 .,...-. 7 10 0 0 0 0 0 +Nature 602, nature N Na Nat Natu BLOCKSTART PAGEIN NEWFONT SAMEFONTSIZE 0 1 INITCAP NODIGIT 0 0 1 0 0 0 0 0 10 4 ,-(). 5 10 0 0 0 0 0 +4. Larson, 4. 4 4. 4. 4. BLOCKSTART PAGEIN NEWFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 10 4 .,...- 6 10 0 0 0 0 0 +reveals tissue-and reveals r re rev reve BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 11 5 --., 4 10 0 0 0 0 0 +2357 (2021). 2357 2 23 235 2357 BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS ALLDIGIT 0 0 0 0 1 0 0 0 11 5 (). 3 10 0 0 0 0 0 +5. Chen, 5. 5 5. 5. 5. BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 11 5 .,..- 5 10 0 0 0 0 0 +and microbes. and a an and and BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 11 6 .,(). 5 10 0 0 0 0 0 +6. Roskams-Hieter, 6. 6 6. 6. 6. BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 11 7 .-,..-- 7 10 0 0 0 0 0 +malignant conditions malignant m ma mal mali BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 11 7 ...,- 5 10 0 0 0 0 0 +(2022). (2022). (2022). ( (2 (20 (202 BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 1 0 0 0 11 8 (). 3 10 0 0 0 0 0 +7. Bryzgunova, 7. 7 7. 7. 7. BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 11 8 .,..,,..,..- 12 10 0 0 0 0 0 +different tumor different d di dif diff BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 11 9 .,-(). 6 10 0 0 0 0 0 +8. Liquori, 8. 8 8. 8. 8. BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 11 9 .,..: 5 10 0 0 0 0 0 +around a around a ar aro arou BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 11 10 -.(),(). 8 10 0 0 0 0 0 +9. Liberzon, 9. 9 9. 9. 9. 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( (h (ht (htt BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 1 0 0 (://../-//---). 15 4 1 1 0 0 1 +Journal canadien journal J Jo Jou Jour BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 1 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 1 no 0 10 0 1 0 0 0 +https://doi.org/10.1177/20543581231183856 https://doi.org/10.1177/20543581231183856 https://doi.org/10.1177/20543581231183856 h ht htt http BLOCKSTART PAGEIN NEWFONT HIGHERFONT 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 1 0 0 1 0 1 ://././ 7 10 0 0 0 0 1 +Canadian Journal canadian C Ca Can Cana BLOCKSTART PAGEIN SAMEFONT HIGHERFONT 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 0 1 no 0 10 0 0 0 0 1 +and Disease and a an and and BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 1 no 0 3 0 0 0 0 1 +Volume 10: volume V Vo Vol Volu BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 1 :- 2 5 0 0 0 0 1 +© The © © © © © BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 1 0 0 0 0 0 0 0 0 1 () 2 6 0 0 0 0 1 +Article reuse article A Ar Art Arti BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 1 : 1 7 0 0 0 0 1 +sagepub.com/journals-permissions sagepub.com/journals-permissions sagepub.com/journals-permissions s sa sag sage BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 0 1 ./- 3 9 0 0 0 0 1 +DOI: 10.1177/20543581231183856 doi: D DO DOI DOI: BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 0 1 :./ 3 9 0 0 0 0 1 +journals.sagepub.com/home/cjk journals.sagepub.com/home/cjk journals.sagepub.com/home/cjk j jo jou jour BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 0 1 ..// 4 8 0 0 0 0 1 +Basic Research basic B Ba Bas Basi BLOCKSTART PAGEIN NEWFONT HIGHERFONT 0 1 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 1 no 0 10 0 1 0 0 0 +1183856C JKXXX10.1177/20543581231183856Canadian 1183856c 1 11 118 1183 BLOCKSTART PAGEIN NEWFONT LOWERFONT 1 0 ALLCAP CONTAINSDIGITS 0 0 0 0 1 0 0 0 0 2 ./ 2 10 0 1 0 0 0 +research-article20232023 research-article20232023 research-article20232023 r re res rese BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 1 NOCAPS CONTAINSDIGITS 0 0 0 0 1 0 0 0 0 2 - 1 10 0 1 0 0 0 +Basic Research basic B Ba Bas Basi BLOCKSTART PAGEIN NEWFONT HIGHERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 2 : 1 8 0 0 0 0 1 +Sequencing in sequencing S Se Seq Sequ BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 0 2 no 0 10 0 0 0 0 1 +Hematuria Syndrome: hematuria H He Hem Hema BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 1 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 0 2 : 1 9 0 0 0 0 1 +Aditi Sharma aditi A Ad Adi Adit BLOCKSTART PAGEIN SAMEFONT LOWERFONT 1 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 0 2 ,.,,, 5 10 0 0 0 0 1 +Pouneh Dokouhaki pouneh P Po Pou Poun BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 1 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 0 2 ,, 2 6 0 0 0 0 1 +Abstract Abstract abstract A Ab Abs Abst BLOCKSTART PAGEIN SAMEFONT LOWERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 2 no 0 0 0 0 0 0 1 +Background: Loin background: B Ba Bac Back BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 2 :() 3 9 0 0 0 0 1 +either unilateral either e ei eit eith BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 2 . 1 10 0 0 0 0 1 +imposes a imposes i im imp impo BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 2 . 1 9 0 0 0 0 1 +to an to t to to to BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 2 ,. 2 9 0 0 0 0 1 +60 years 60 6 60 60 60 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS ALLDIGIT 0 0 0 0 0 0 0 0 0 2 ,. 2 9 0 0 0 0 1 +Objective: To objective: O Ob Obj Obje BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 2 :. 2 7 0 0 0 0 1 +Methods: In methods: M Me Met Meth BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 2 :-,- 4 9 0 0 0 0 1 +will be will w wi wil will BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS NODIGIT 0 1 1 0 0 0 0 0 0 2 . 1 8 0 0 0 0 1 +6000 System 6000 6 60 600 6000 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS ALLDIGIT 0 0 0 0 0 0 0 0 0 2 ( 1 9 0 0 0 0 1 +genes in: genes g ge gen gene BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 0 2 :[][]),(: 9 9 0 0 0 0 1 +transduction [n transduction t tr tra tran BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 0 2 [],[],[],[]). 13 9 0 0 0 0 1 +examine identified examine e ex exa exam BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 0 2 -. 2 8 0 0 0 0 1 +Conclusions: This conclusions: C Co Con Conc BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 0 2 : 1 9 0 0 0 0 1 +LPHS. 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In ants. a an ant ants BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 0 ., 2 9 1 1 0 0 1 +the other the t th the the BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 0 . 1 8 1 1 0 0 1 +Systematic Review systematic S Sy Sys Syst BLOCKSTART PAGEIN NEWFONT HIGHERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 5 no 0 10 0 0 0 0 1 +Studies Studies studies S St Stu Stud BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 5 no 0 2 0 0 0 0 1 +We searched we W We We We BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 5 ,, 2 8 0 0 0 0 1 +databases to databases d da dat data BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 5 (: 2 10 0 0 0 0 1 +"loin pain "loin " "l "lo "loi BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 3 5 """-"" 6 9 0 0 0 0 1 +pain/haematuria" OR pain/haematuria" p pa pai pain BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 3 6 /"""). 6 8 1 1 0 0 1 +was generated was w wa was was BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 6 ,. 2 8 1 1 0 0 1 +total of total t to tot tota BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 6 . 1 9 1 1 0 0 1 +or articles or o or or or BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 6 (). 3 8 1 1 0 0 1 +articles included, articles a ar art arti BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 6 ,, 2 8 1 1 0 0 1 +68% of 68% 6 68 68% 68% BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 6 no 0 8 1 1 0 0 1 +United States united U Un Uni Unit BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 6 . 1 8 1 1 0 0 1 +published so published p pu pub publ BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 6 no 0 8 1 1 0 0 1 +using oral using u us usi usin BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 6 / 1 8 1 1 0 0 1 +strategies such strategies s st str stra BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 6 -,- 3 9 1 1 0 0 1 +denervation, 18-20 denervation, d de den dene BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 3 6 ,-,-- 5 9 1 1 0 0 1 +tion, 24-26 tion, t ti tio tion BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 3 6 ,--,- 5 10 1 1 0 0 1 +studies looking studies s st stu stud BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 6 () 2 9 1 1 0 0 1 +or its or o or or or BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 6 , 1 9 1 1 0 0 1 +ancestries, or ancestries, a an anc ance BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 6 ,,()- 5 9 1 1 0 0 1 +ants in ants a an ant ants BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 6 , 1 8 1 1 0 0 1 +GBM, or gbm, G GB GBM GBM, BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 6 ,. 2 3 1 1 0 0 1 +Research Objectives research R Re Res Rese BLOCKSTART PAGEIN NEWFONT HIGHERFONT 1 0 INITCAP NODIGIT 0 1 1 0 0 0 0 0 3 10 no 0 10 0 0 0 0 1 +The specific the T Th The The BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 10 : 1 10 0 0 0 0 1 +Objective 1: objective O Ob Obj Obje BLOCKSTART PAGEIN NEWFONT SAMEFONTSIZE 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 10 : 1 10 0 0 0 0 1 +18 genes 18 1 18 18 18 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS ALLDIGIT 0 0 0 0 0 0 0 0 3 10 (). 3 8 0 0 0 0 1 +Objective 2: objective O Ob Obj Obje BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 10 : 1 10 0 0 0 0 1 +89 genes 89 8 89 89 89 BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 1 0 NOCAPS ALLDIGIT 0 0 0 0 0 0 0 0 3 10 (). 3 9 0 0 0 0 1 +Methods Methods methods M Me Met Meth BLOCKSTART PAGEIN NEWFONT HIGHERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 11 no 0 10 0 0 0 0 1 +We plan we W We We We BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 11 -, 2 10 0 0 0 0 1 +of decoding of o of of of BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 3 11 . 1 9 0 0 0 0 1 +Figure 1. figure F Fi Fig Figu BLOCKSTART PAGEEND NEWFONT LOWERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 11 .. 2 10 1 1 0 0 1 +4 4 4 4 4 4 4 BLOCKSTART PAGESTART NEWFONT HIGHERFONT 0 0 NOCAPS ALLDIGIT 1 0 0 0 0 0 0 0 3 0 no 0 10 0 1 0 0 0 +Canadian Journal canadian C Ca Can Cana BLOCKSTART PAGEIN NEWFONT SAMEFONTSIZE 0 1 INITCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 10 0 1 1 0 0 +Table 1. table T Ta Tab Tabl BLOCKSTART PAGEIN NEWFONT LOWERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 0 .. 2 10 0 1 0 0 1 +Gene name gene G Ge Gen Gene BLOCKSTART PAGEIN NEWFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 1 0 0 0 0 0 0 3 0 no 0 5 0 1 0 0 1 +Chromosome number chromosome C Ch Chr Chro BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 0 no 0 10 0 1 0 0 1 +Inheritance a inheritance I In Inh Inhe BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 0 no 0 7 0 1 0 0 1 +Gene annotation gene G Ge Gen Gene BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 1 0 0 0 0 0 0 3 0 no 0 8 0 1 0 0 1 +Glomerular basement glomerular G Gl Glo Glom BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 9 0 0 0 0 1 +LAMB2 LAMB2 lamb2 L LA LAM LAMB BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +chr3 chr3 chr3 c ch chr chr3 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +AR AR ar A AR AR AR BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 0 0 0 0 0 1 +Laminin subunit laminin L La Lam Lami BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 5 0 0 0 0 1 +LAMA5 LAMA5 lama5 L LA LAM LAMA BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +chr20 chr20 chr20 c ch chr chr2 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +AR AR ar A AR AR AR BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 0 0 0 0 0 1 +Laminin subunit laminin L La Lam Lami BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 5 0 0 0 0 1 +ITGA3 ITGA3 itga3 I IT ITG ITGA BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +chr17 chr17 chr17 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +AR AR ar A AR AR AR BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 0 0 0 0 0 1 +Integrin subunit integrin I In Int Inte BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 5 0 0 0 0 1 +COL4A3/4/5 COL4A3/4/5 col4a3/4/5 C CO COL COL4 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 // 2 2 0 0 0 0 1 +chr2/chr2/chrX chr2/chr2/chrX chr2/chr2/chrx c ch chr chr2 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 // 2 3 0 0 0 0 1 +AR, AD/AR, ar, A AR AR, AR, BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 ,/,/ 4 4 0 0 0 0 1 +Collagen type collagen C Co Col Coll BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 // 2 7 0 0 0 0 1 +GPC5 GPC5 gpc5 G GP GPC GPC5 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +chr13 chr13 chr13 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +Glypican 5 glypican G Gl Gly Glyp BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 2 0 0 0 0 1 +CD151 CD151 cd151 C CD CD1 CD15 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +chr11 chr11 chr11 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +CD151 molecule cd151 C CD CD1 CD15 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 - 1 10 0 0 0 0 1 +Endothelial cells endothelial E En End Endo BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 4 0 0 0 0 1 +CFH CFH cfh C CF CFH CFH BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 0 0 0 0 0 1 +chr1 chr1 chr1 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +AR, AD ar, A AR AR, AR, BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 , 1 1 0 0 0 0 1 +Complement factor complement C Co Com Comp BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 0 no 0 4 0 0 0 0 1 +CFB CFB cfb C CF CFB CFB BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 0 0 0 0 0 1 +chr6 chr6 chr6 c ch chr chr6 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +AR, AD, ar, A AR AR, AR, BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 ,, 2 2 0 0 0 0 1 +Complement factor complement C Co Com Comp BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 0 no 0 4 0 0 0 0 1 +CFI CFI cfi C CF CFI CFI BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 0 0 0 0 0 1 +chr4 chr4 chr4 c ch chr chr4 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +AR, AD ar, A AR AR, AR, BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 , 1 1 0 0 0 0 1 +Complement factor complement C Co Com Comp BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 0 no 0 4 0 0 0 0 1 +C3 C3 c3 C C3 C3 C3 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 0 0 0 0 0 1 +chr19 chr19 chr19 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +AR, AD ar, A AR AR, AR, BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 , 1 1 0 0 0 0 1 +Complement C3 complement C Co Com Comp BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 0 no 0 3 0 0 0 0 1 +MCP MCP mcp M MC MCP MCP BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 0 0 0 0 0 1 +chr1 chr1 chr1 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +AR, AD ar, A AR AR, AR, BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 , 1 1 0 0 0 0 1 +Membrane cofactor membrane M Me Mem Memb BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 0 no 0 5 0 0 0 0 1 +THBD THBD thbd T TH THB THBD BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +chr20 chr20 chr20 c ch chr chr2 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +AD AD ad A AD AD AD BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 1 0 0 0 0 0 3 0 no 0 0 0 0 0 0 1 +Thrombomodulin Thrombomodulin thrombomodulin T Th Thr Thro BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 3 0 0 0 0 1 +PLG PLG plg P PL PLG PLG BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 0 0 0 0 0 1 +chr6 chr6 chr6 c ch chr chr6 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +AR, AD ar, A AR AR, AR, BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 , 1 1 0 0 0 0 1 +Plasminogen Plasminogen plasminogen P Pl Pla Plas BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 2 0 0 0 0 1 +DGKE DGKE dgke D DG DGK DGKE BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +chr17 chr17 chr17 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +AR AR ar A AR AR AR BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 0 0 0 0 0 1 +Diacylglycerol kinase diacylglycerol D Di Dia Diac BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 6 0 0 0 0 1 +INF2 INF2 inf2 I IN INF INF2 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +chr14 chr14 chr14 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +AD AD ad A AD AD AD BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 1 0 0 0 0 0 3 0 no 0 0 0 0 0 0 1 +Inverted formin inverted I In Inv Inve BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 0 no 0 4 0 0 0 0 1 +MMACHC MMACHC mmachc M MM MMA MMAC BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +chr1 chr1 chr1 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 3 0 no 0 1 0 0 0 0 1 +AR AR ar A AR AR AR BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 3 0 no 0 0 0 0 0 0 1 +Metabolism of metabolism M Me Met Meta BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 3 0 no 0 8 0 0 0 0 1 +a a a a a a a BLOCKSTART PAGEIN SAMEFONT LOWERFONT 0 0 NOCAPS NODIGIT 1 0 1 0 0 0 0 0 4 4 no 0 10 0 0 0 0 1 +Inheritance based inheritance I In Inh Inhe BLOCKSTART PAGEIN SAMEFONT HIGHERFONT 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 4 :,;,;,;,;,;, 12 10 0 0 0 0 1 +X linked x X X X X BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 1 0 0 0 0 0 0 0 4 4 ;, 2 2 0 0 0 0 1 +Table 2. table T Ta Tab Tabl BLOCKSTART PAGEIN NEWFONT HIGHERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 .. 2 10 0 0 0 0 1 +Gene name gene G Ge Gen Gene BLOCKSTART PAGEIN NEWFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 1 0 0 0 0 0 0 4 5 no 0 5 0 0 0 0 1 +Chromosome number chromosome C Ch Chr Chro BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 no 0 10 0 0 0 0 1 +Inheritance a inheritance I In Inh Inhe BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 no 0 7 0 0 0 0 1 +Gene annotation gene G Ge Gen Gene BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 1 0 0 0 0 0 0 4 5 no 0 8 0 0 0 0 1 +Pain transduction pain P Pa Pai Pain BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 no 0 3 0 0 0 0 1 +TRPV1/2/3/4 TRPV1/2/3/4 trpv1/2/3/4 T TR TRP TRPV BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 /// 3 2 0 0 0 0 1 +chr17/chr17/chr17/chr12 TRPV3/4: chr17/chr17/chr17/chr12 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 ////: 5 6 0 0 0 0 1 +Transient receptor transient T Tr Tra Tran BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 no 0 10 0 0 0 0 1 +V member v V V V V BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 1 0 0 0 0 0 0 0 4 5 /// 3 3 0 0 0 0 1 +P2RX3 P2RX3 p2rx3 P P2 P2R P2RX BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 1 0 0 0 0 1 +chr11 chr11 chr11 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 1 0 0 0 0 1 +P2X purinergic p2x P P2 P2X P2X BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 4 0 0 0 0 1 +TRPM8 TRPM8 trpm8 T TR TRP TRPM BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 1 0 0 0 0 1 +chr2 chr2 chr2 c ch chr chr2 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 0 0 0 0 0 1 +Transient receptor transient T Tr Tra Tran BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 no 0 10 0 0 0 0 1 +M member m M M M M BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 1 0 0 0 0 0 0 0 4 5 no 0 2 0 0 0 0 1 +TRPA1 TRPA1 trpa1 T TR TRP TRPA BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 1 0 0 0 0 1 +chr8 chr8 chr8 c ch chr chr8 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 0 0 0 0 0 1 +AD AD ad A AD AD AD BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 no 0 0 0 0 0 0 1 +Transient receptor transient T Tr Tra Tran BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 no 0 10 0 0 0 0 1 +A member a A A A A BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 1 0 1 0 0 0 0 0 4 5 no 0 2 0 0 0 0 1 +P2RY12 P2RY12 p2ry12 P P2 P2R P2RY BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 1 0 0 0 0 1 +chr3 chr3 chr3 c ch chr chr3 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 0 0 0 0 0 1 +AR AR ar A AR AR AR BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 5 no 0 0 0 0 0 0 1 +Purinergic receptor purinergic P Pu Pur Puri BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 4 5 no 0 4 0 0 0 0 1 +BDKRB1/2 BDKRB1/2 bdkrb1/2 B BD BDK BDKR BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 / 1 1 0 0 0 0 1 +chr14/chr14 chr14/chr14 chr14/chr14 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 / 1 2 0 0 0 0 1 +Bradykinin receptor bradykinin B Br Bra Brad BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 4 5 / 1 4 0 0 0 0 1 +HTR3a HTR3a htr3a H HT HTR HTR3 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 1 0 0 0 0 1 +chr11 chr11 chr11 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 1 0 0 0 0 1 +5-Hydroxytryptamine receptor 5-hydroxytryptamine 5 5- 5-H 5-Hy BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 - 1 5 0 0 0 0 1 +ACCN1/2 ACCN1/2 accn1/2 A AC ACC ACCN BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 / 1 1 0 0 0 0 1 +chr17/chr12 chr17/chr12 chr17/chr12 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 / 1 2 0 0 0 0 1 +Acid sensing acid A Ac Aci Acid BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 / 1 6 0 0 0 0 1 +TRPC/P TRPC/P trpc/p T TR TRP TRPC BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 5 / 1 1 0 0 0 0 1 +chr13 chr13 chr13 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 1 0 0 0 0 1 +Transient receptor transient T Tr Tra Tran BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 no 0 7 0 0 0 0 1 +Pain conduction pain P Pa Pai Pain BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 no 0 2 0 0 0 0 1 +SCN10A SCN10A scn10a S SC SCN SCN1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 1 0 0 0 0 1 +chr3 chr3 chr3 c ch chr chr3 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 0 0 0 0 0 1 +AD AD ad A AD AD AD BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 no 0 0 0 0 0 0 1 +Sodium voltage-gated sodium S So Sod Sodi BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 - 1 8 0 0 0 0 1 +SCN11A SCN11A scn11a S SC SCN SCN1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 1 0 0 0 0 1 +chr3 chr3 chr3 c ch chr chr3 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 0 0 0 0 0 1 +AD AD ad A AD AD AD BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 no 0 0 0 0 0 0 1 +Sodium voltage-gated sodium S So Sod Sodi BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 - 1 8 0 0 0 0 1 +SCN1,3,8A SCN1,3,8A scn1,3,8a S SC SCN SCN1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 ,, 2 1 0 0 0 0 1 +chr2/chr2/chr12 chr2/chr2/chr12 chr2/chr2/chr12 c ch chr chr2 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 // 2 2 0 0 0 0 1 +AD/AD/AD AD/AD/AD ad/ad/ad A AD AD/ AD/A BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 5 // 2 1 0 0 0 0 1 +Sodium channel sodium S So Sod Sodi BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 // 2 6 0 0 0 0 1 +SCN9A SCN9A scn9a S SC SCN SCN9 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 1 0 0 0 0 1 +chr2 chr2 chr2 c ch chr chr2 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 0 0 0 0 0 1 +AR, AD ar, A AR AR, AR, BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 5 , 1 1 0 0 0 0 1 +Sodium voltage-gated sodium S So Sod Sodi BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 - 1 8 0 0 0 0 1 +KCNQ KCNQ kcnq K KC KCN KCNQ BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 5 no 0 0 0 0 0 0 1 +chr11 chr11 chr11 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 1 0 0 0 0 1 +AR, AD ar, A AR AR, AR, BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 5 , 1 1 0 0 0 0 1 +Potassium voltage-gated potassium P Po Pot Pota BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 - 1 8 0 0 0 0 1 +Pain synaptic pain P Pa Pai Pain BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 no 0 5 0 0 0 0 1 +NR1, 2 nr1, N NR NR1 NR1, BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 , 1 1 0 0 0 0 1 +chr9/ chr9/ chr9/ c ch chr chr9 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 / 1 1 0 0 0 0 1 +AR, AD ar, A AR AR, AR, BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 5 , 1 1 0 0 0 0 1 +Nuclear receptor nuclear N Nu Nuc Nucl BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 5 / 1 5 0 0 0 0 1 +GRIA1-4 GRIA1-4 gria1-4 G GR GRI GRIA BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 - 1 1 0 0 0 0 1 +chr5/chr4/chrX/chr11 chr5/chr4/chrX/chr11 chr5/chr4/chrx/chr11 c ch chr chr5 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 /// 3 3 0 0 0 0 1 +AR, AD/ ar, A AR AR, AR, BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 5 ,/// 4 3 0 0 0 0 1 +AMPA receptor ampa A AM AMP AMPA BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 5 - 1 3 0 0 0 0 1 +NK1R NK1R nk1r N NK NK1 NK1R BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 0 0 0 0 0 1 +chr2 chr2 chr2 c ch chr chr2 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 no 0 0 0 0 0 0 1 +Tachykinin receptor tachykinin T Ta Tac Tach BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 4 5 no 0 4 0 0 0 0 1 +CACNA1A-I, S cacna1a-i, C CA CAC CACN BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 -, 2 2 0 0 0 0 1 +chr19/chr9/chr12/chr3/ chr19/chr9/chr12/chr3/ chr19/chr9/chr12/chr3/ c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 //// 4 4 0 0 0 0 1 +chr1/chrX/chr17/chr16/ chr1/chrX/chr17/chr16/ chr1/chrx/chr17/chr16/ c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 //// 4 4 0 0 0 0 1 +chr22/chr1 chr22/chr1 chr22/chr1 c ch chr chr2 BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 5 / 1 2 0 0 0 0 1 +AD/ AR/ ad/ A AD AD/ AD/ BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 10 ///,// 6 10 0 1 0 0 1 +XL, XLR/ xl, X XL XL, XL, BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 10 ,//// 5 9 0 1 0 0 1 +R, AD r, R R, R, R, BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 10 , 1 2 0 1 0 0 1 +Calcium voltage-gated calcium C Ca Cal Calc BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 11 -- 2 10 0 1 0 0 1 +CACNA2D1 CACNA2D1 cacna2d1 C CA CAC CACN BLOCKSTART PAGEIN NEWFONT SAMEFONTSIZE 0 1 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 11 no 0 1 0 1 0 0 1 +chr7 chr7 chr7 c ch chr chr7 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 11 no 0 0 0 1 0 0 1 +AR AR ar A AR AR AR BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 11 no 0 0 0 1 0 0 1 +Calcium voltage-gated calcium C Ca Cal Calc BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 11 - 1 10 0 1 0 0 1 +delta 1 delta d de del delt BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS NODIGIT 0 1 1 0 0 0 0 0 4 11 no 0 1 0 1 0 0 1 +(continued) (continued) (continued) ( (c (co (con BLOCKSTART PAGEEND SAMEFONT SAMEFONTSIZE 0 1 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 4 11 () 2 10 0 1 0 0 1 +Sharma et sharma S Sh Sha Shar BLOCKSTART PAGESTART SAMEFONT HIGHERFONT 0 1 INITCAP NODIGIT 0 1 0 0 0 0 0 0 4 0 no 0 10 0 1 1 0 0 +5 5 5 5 5 5 5 BLOCKSTART PAGEIN NEWFONT SAMEFONTSIZE 0 0 NOCAPS ALLDIGIT 1 0 0 0 0 0 0 0 4 0 no 0 10 0 1 0 0 0 +Gene name gene G Ge Gen Gene BLOCKSTART PAGEIN SAMEFONT LOWERFONT 0 0 INITCAP NODIGIT 0 1 0 0 0 0 0 0 4 0 no 0 5 0 1 0 0 1 +Chromosome number chromosome C Ch Chr Chro BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 0 no 0 10 0 1 0 0 1 +Inheritance a inheritance I In Inh Inhe BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 0 no 0 7 0 1 0 0 1 +Gene annotation gene G Ge Gen Gene BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 1 0 0 0 0 0 0 4 0 no 0 8 0 1 0 0 1 +Pain modulation pain P Pa Pai Pain BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 0 no 0 4 0 0 0 0 1 +BDNF BDNF bdnf B BD BDN BDNF BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 0 no 0 1 0 0 0 0 1 +chr11 chr11 chr11 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 0 no 0 1 0 0 0 0 1 +Brain-derived neurotrophic brain-derived B Br Bra Brai BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 0 - 1 9 0 0 0 0 1 +OPRD1/M1/K1 OPRD1/M1/K1 oprd1/m1/k1 O OP OPR OPRD BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 0 // 2 3 0 0 0 0 1 +chr1/ chr6/ chr1/ c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 0 // 2 4 0 0 0 0 1 +Opioid receptor opioid O Op Opi Opio BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 4 0 // 2 10 0 0 0 0 1 +CNR1 CNR1 cnr1 C CN CNR CNR1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 0 no 0 1 0 0 0 0 1 +chr6 chr6 chr6 c ch chr chr6 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 0 no 0 1 0 0 0 0 1 +Cannabinoid receptor cannabinoid C Ca Can Cann BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 4 0 no 0 6 0 0 0 0 1 +GABRs (GABRD/E/ gabrs G GA GAB GABR BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 4 0 (// 3 4 0 0 0 0 1 +P/Q/A1-A6/BR1- P/Q/A1-A6/BR1- p/q/a1-a6/br1- P P/ P/Q P/Q/ BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 0 //-/- 5 3 0 0 0 0 1 +BR3/G1-G3/R1-R3 BR3/G1-G3/R1-R3 br3/g1-g3/r1-r3 B BR BR3 BR3/ BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 0 /-/- 4 4 0 0 0 0 1 +chr1/chrX/chr5/chrX/chr5/ chr1/chrX/chr5/chrX/chr5/ chr1/chrx/chr5/chrx/chr5/ c ch chr chr1 BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 1 ///// 5 10 0 0 0 0 1 +chr4/chrX/chr4/chr15/ chr4/chrX/chr4/chr15/ chr4/chrx/chr4/chr15/ c ch chr chr4 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 1 //// 4 8 0 0 0 0 1 +chr5/chr4/chr5/chr15/ chr5/chr4/chr5/chr15/ chr5/chr4/chr5/chr15/ c ch chr chr5 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 1 //// 4 8 0 0 0 0 1 +chr4/chr5/chr15/chr6/ chr4/chr5/chr15/chr6/ chr4/chr5/chr15/chr6/ c ch chr chr4 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 1 //// 4 8 0 0 0 0 1 +chr6/chr3 chr6/chr3 chr6/chr3 c ch chr chr6 BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 1 / 1 3 0 0 0 0 1 +AD/ M/ ad/ A AD AD/ AD/ BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 1 /////,/ 7 10 0 0 0 0 1 +XL/ M/ xl/ X XL XL/ XL/ BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 1 ////// 6 8 0 0 0 0 1 +AD/ M/ ad/ A AD AD/ AD/ BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 1 ////// 6 8 0 0 0 0 1 +Gamma-aminobutyric acid gamma-aminobutyric G Ga Gam Gamm BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 4 2 - 1 10 0 0 0 0 1 +TNF TNF tnf T TN TNF TNF BLOCKSTART PAGEIN NEWFONT SAMEFONTSIZE 0 1 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 2 no 0 0 0 0 0 0 1 +chr6 chr6 chr6 c ch chr chr6 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 no 0 1 0 0 0 0 1 +AD AD ad A AD AD AD BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP NODIGIT 0 0 1 0 0 0 0 0 4 2 no 0 0 0 0 0 0 1 +Tumor necrosis tumor T Tu Tum Tumo BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 2 no 0 4 0 0 0 0 1 +PLA2G2A PLA2G2A pla2g2a P PL PLA PLA2 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 no 0 1 0 0 0 0 1 +chr1 chr1 chr1 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 no 0 1 0 0 0 0 1 +AD, SMu ad, A AD AD, AD, BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP NODIGIT 0 0 1 0 0 0 0 0 4 2 , 1 1 0 0 0 0 1 +Phospholipase A2 phospholipase P Ph Pho Phos BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 INITCAP NODIGIT 0 0 0 0 0 0 0 0 4 2 no 0 3 0 0 0 0 1 +IL1/6/12/18 IL1/6/12/18 il1/6/12/18 I IL IL1 IL1/ BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 /// 3 2 0 0 0 0 1 +chr2/chr7/chr3/chr11 chr2/chr7/chr3/chr11 chr2/chr7/chr3/chr11 c ch chr chr2 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 /// 3 4 0 0 0 0 1 +M/ Mu, m/ M M/ M/ M/ BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 2 /,,,///// 9 10 0 0 0 0 1 +COX2 COX2 cox2 C CO COX COX2 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 no 0 1 0 0 0 0 1 +chrMT chrMT chrmt c ch chr chrM BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 4 2 no 0 1 0 0 0 0 1 +Cytochrome C cytochrome C Cy Cyt Cyto BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 INITCAP NODIGIT 0 0 0 0 0 0 0 0 4 2 no 0 6 0 0 0 0 1 +NTRK1 NTRK1 ntrk1 N NT NTR NTRK BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 no 0 1 0 0 0 0 1 +chr1 chr1 chr1 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 no 0 1 0 0 0 0 1 +AR AR ar A AR AR AR BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 2 no 0 0 0 0 0 0 1 +High affinity high H Hi Hig High BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 INITCAP NODIGIT 0 1 1 0 0 0 0 0 4 2 no 0 9 0 0 0 0 1 +NGF NGF ngf N NG NGF NGF BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 2 no 0 0 0 0 0 0 1 +chr1 chr1 chr1 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 no 0 1 0 0 0 0 1 +AR AR ar A AR AR AR BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 2 no 0 0 0 0 0 0 1 +Beta-nerve growth beta-nerve B Be Bet Beta BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 2 - 1 5 0 0 0 0 1 +GDNF GDNF gdnf G GD GDN GDNF BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 2 no 0 1 0 0 0 0 1 +chr5 chr5 chr5 c ch chr chr5 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 no 0 1 0 0 0 0 1 +AD AD ad A AD AD AD BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP NODIGIT 0 0 1 0 0 0 0 0 4 2 no 0 0 0 0 0 0 1 +Glial cell glial G Gl Gli Glia BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 INITCAP NODIGIT 0 0 0 0 0 0 0 0 4 2 - 1 9 0 0 0 0 1 +LIF LIF lif L LI LIF LIF BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 2 no 0 0 0 0 0 0 1 +chr22 chr22 chr22 c ch chr chr2 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 no 0 1 0 0 0 0 1 +AD AD ad A AD AD AD BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP NODIGIT 0 0 1 0 0 0 0 0 4 2 no 0 0 0 0 0 0 1 +Leukemia inhibitory leukemia L Le Leu Leuk BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 INITCAP NODIGIT 0 0 0 0 0 0 0 0 4 2 no 0 5 0 0 0 0 1 +CCL2 CCL2 ccl2 C CC CCL CCL2 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 no 0 1 0 0 0 0 1 +chr17 chr17 chr17 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 no 0 1 0 0 0 0 1 +C-C motif c-c C C- C-C C-C BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP NODIGIT 0 0 0 0 0 0 0 0 4 2 - 1 4 0 0 0 0 1 +CNR2 CNR2 cnr2 C CN CNR CNR2 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 no 0 1 0 0 0 0 1 +chr1 chr1 chr1 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 no 0 1 0 0 0 0 1 +Cannabinoid receptor cannabinoid C Ca Can Cann BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 INITCAP NODIGIT 0 0 0 0 0 0 0 0 4 2 no 0 4 0 0 0 0 1 +TLR2/4 TLR2/4 tlr2/4 T TL TLR TLR2 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 / 1 1 0 0 0 0 1 +chr4/chr9 chr4/chr9 chr4/chr9 c ch chr chr4 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 / 1 2 0 0 0 0 1 +AD, SMu/ ad, A AD AD, AD, BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP NODIGIT 0 0 1 0 0 0 0 0 4 2 ,/ 2 2 0 0 0 0 1 +Toll-like receptor toll-like T To Tol Toll BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 INITCAP NODIGIT 0 0 1 0 0 0 0 0 4 2 -/ 2 4 0 0 0 0 1 +P2RX4/7 P2RX4/7 p2rx4/7 P P2 P2R P2RX BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 / 1 1 0 0 0 0 1 +chr12/chr12 chr12/chr12 chr12/chr12 c ch chr chr1 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 / 1 2 0 0 0 0 1 +P2X purinoceptor p2x P P2 P2X P2X BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 / 1 4 0 0 0 0 1 +CX3CR1 CX3CR1 cx3cr1 C CX CX3 CX3C BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 no 0 1 0 0 0 0 1 +chr3 chr3 chr3 c ch chr chr3 BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 1 NOCAPS CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 no 0 1 0 0 0 0 1 +CX3C chemokine cx3c C CX CX3 CX3C BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 1 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 4 2 no 0 5 0 0 0 0 1 +a a a a a a a BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 NOCAPS NODIGIT 1 0 1 0 0 0 0 0 5 4 no 0 10 0 1 0 0 1 +Inheritance based inheritance I In Inh Inhe BLOCKSTART PAGEIN SAMEFONT HIGHERFONT 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 5 4 :,;,;,;,;,;, 12 10 0 1 0 0 1 +X linked x X X X X BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP NODIGIT 1 0 0 0 0 0 0 0 5 4 ;,. 3 4 0 1 0 0 1 +Table 2. table T Ta Tab Tabl BLOCKSTART PAGEIN NEWFONT HIGHERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 5 5 .() 3 10 0 1 0 0 1 +been approved been b be bee been BLOCKSTART PAGEIN NEWFONT HIGHERFONT 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 5 5 no 0 10 0 1 0 0 1 +Research Ethics research R Re Res Rese BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 1 1 0 0 0 0 0 5 5 (--). 5 6 0 1 0 0 1 +Design Design design D De Des Desi BLOCKSTART PAGEIN NEWFONT HIGHERFONT 1 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 5 5 no 0 10 0 1 0 0 1 +Number of number N Nu Num Numb BLOCKSTART PAGEIN NEWFONT SAMEFONTSIZE 0 1 INITCAP NODIGIT 0 0 1 0 0 0 0 0 5 5 no 0 10 0 0 0 0 1 +Twenty-four LPHS twenty-four T Tw Twe Twen BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 5 5 - 1 9 0 0 0 0 1 +(run by (run ( (r (ru (run BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 5 5 () 2 9 0 0 0 0 1 +study coordinator, study s st stu stud BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 5 5 , 1 9 0 0 0 0 1 +study. Recruitment study. s st stu stud BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 5 5 . 1 9 0 0 0 0 1 +(parents/siblings/cousins/children) is (parents/siblings/cousins/children) ( (p (pa (par BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 5 5 (///). 6 9 0 0 0 0 1 +first study first f fi fir firs BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 5 5 (, 2 10 0 0 0 0 1 +17) with 17) 1 17 17) 17) BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 5 5 )(). 4 9 0 0 0 0 1 +addition, 6 addition, a ad add addi BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 5 5 ,(:,,,,,) 9 10 0 0 0 0 1 +had a had h ha had had BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 5 5 no 0 9 0 0 0 0 1 +(Figure 2). (figure ( (F (Fi (Fig BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 5 5 (). 3 1 0 0 0 0 1 +Duration of duration D Du Dur Dura BLOCKSTART PAGEIN NEWFONT HIGHERFONT 0 1 INITCAP NODIGIT 0 0 1 0 0 0 0 0 5 7 no 0 10 0 0 0 0 1 +The study the T Th The The BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 5 7 (-). 4 10 0 0 0 0 1 +Participant Selection participant P Pa Par Part BLOCKSTART PAGEIN NEWFONT HIGHERFONT 0 1 INITCAP NODIGIT 0 0 1 0 0 0 0 0 5 8 no 0 10 0 0 0 0 1 +The LPHS the T Th The The BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 INITCAP NODIGIT 0 0 1 0 0 0 0 0 5 8 () 2 9 0 0 0 0 1 +and FG and a an and and BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 5 8 (),- 4 9 0 0 0 0 1 +teria defined teria t te ter teri BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 5 8 - 1 9 0 0 0 0 1 +bers that bers b be ber bers BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 5 8 no 0 10 0 0 0 0 1 +the written the t th the the BLOCKSTART PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 5 8 - 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Little 1. 1 1. 1. 1. BLOCKSTART PAGEIN NEWFONT LOWERFONT 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 8 5 .,,. 4 10 0 0 0 0 1 +and haematuria and a an and and BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 8 5 no 0 9 0 0 0 0 1 +arteries. Q arteries. a ar art arte BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 8 5 ..;():- 7 6 0 0 0 0 1 +2. Vakili 2. 2 2. 2. 2. BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 8 5 .,,.- 5 9 0 0 0 0 1 +drome. Am drome. d dr dro drom BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 8 5 ..;():-.:./. 12 9 0 0 0 0 1 +ajkd.2014.01.439 ajkd.2014.01.439 ajkd.2014.01.439 a aj ajk ajkd BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS CONTAINSDIGITS 0 0 0 0 1 0 0 0 8 5 ... 3 2 0 0 0 0 1 +3. Aber 3. 3 3. 3. 3. BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 8 5 .,. 3 9 0 0 0 0 1 +the loin the t th the the BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 1 0 0 0 0 0 8 5 /..;():- 8 9 0 0 0 0 1 +615. doi:10.1111/j.1464-410x.1982.tb13607.x 615. 6 61 615 615. BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 8 5 .:./.-... 9 6 0 0 0 0 1 +4. Spetie 4. 4 4. 4. 4. BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 ALLCAP CONTAINSDIGITS 0 0 0 0 0 0 0 0 8 5 .,,,.- 6 9 0 0 0 0 1 +esis of esis e es esi esis BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 8 5 -. 2 9 0 0 0 0 1 +Dis. 2006;47(3):419-427. dis. D Di Dis Dis. BLOCKIN PAGEIN SAMEFONT SAMEFONTSIZE 0 0 INITCAP NODIGIT 0 0 0 0 0 0 0 0 8 5 .;():-.:./.... 14 8 0 0 0 0 1 +5. Cox 5. 5 5. 5. 5. 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Published html. h ht htm html BLOCKEND PAGEIN SAMEFONT SAMEFONTSIZE 0 0 NOCAPS NODIGIT 0 0 0 0 0 0 0 0 11 6 .,.,. 5 8 0 1 0 0 1 + diff --git a/grobid-trainer/resources/dataset/segmentation/corpus/tei/pub.1158465915.training.segmentation.tei.xml b/grobid-trainer/resources/dataset/segmentation/corpus/tei/pub.1158465915.training.segmentation.tei.xml new file mode 100644 index 0000000000..2889ebaaf7 --- /dev/null +++ b/grobid-trainer/resources/dataset/segmentation/corpus/tei/pub.1158465915.training.segmentation.tei.xml @@ -0,0 +1,23 @@ + + + + + + + Title Patient-specific alterations in blood plasma cfRNA profiles enable accurate classification of cancer patients and controls Authors Annelien Morlion (1,2), Philippe Decruyenaere (1,2,3), Kathleen Schoofs (1,2,4), Jasper Anckaert (1,2), Justine Nuytens (1,2), Eveline Vanden Eynde (1,2), Kimberly Verniers (1,2), Celine Everaert (4), Fritz Offner (3), Jo Van Dorpe (5), Jo Vandesompele° (1,2), Pieter Mestdagh°* (1,2) (1) Department of Biomolecular Medicine, Ghent University, Ghent, Belgium (2) OncoRNALab, Cancer Research Institute Ghent (CRIG), Ghent, Belgium (3) Department of Hematology, Ghent University Hospital, Ghent, Belgium (4) TOBI Lab, Center for Medical Biotechnology, VIB-UGent, Zwijnaarde, Belgium (5) Department of Pathology, Ghent University Hospital, Ghent, Belgium °Both authors contributed equally to this work *Corresponding author Author contributions Conceptualization: AM, CE, JV, PM Data Curation: AM Funding acquisition: AM, JV, JVD, PM Software: AM, JA Validation: AM Formal analysis: AM Investigation: EV, JN, KV Methodology: AM, JV, PM Project administration: AM, JV, PM Resources: FO, JV, JVD, KS, PD, PM Supervision: JV, PM Visualization: AM Writing -original draft: AM, JV, PM Writing -review & editing: all authors Abstract Circulating nucleic acids in blood plasma form an attractive resource to study human health and disease. Here, we applied mRNA capture sequencing of blood plasma cell-free RNA from 266 cancer patients and cancer-free controls (discovery n=208, 25 cancer types; validation n=58, 3 types). We observed cancer-type specific as well as pan-cancer alterations in cell-free transcriptomes compared to controls. Differentially abundant RNAs were heterogenous among patients and among cohorts, hampering identification of robust cancer biomarkers. Therefore, we developed a novel method that compares each individual cancer patient to a reference control population to identify so-called biomarker tail genes. These biomarker tail genes discriminate ovarian, prostate, and uterine cancer patients from controls with very high accuracy (AUC = 0.980). Our results were confirmed in additional cohorts of 65 plasma donors (2 lymphoma types) and 24 urine donors (bladder cancer). Together, our findings demonstrate heterogeneity in cell-free RNA alterations among cancer patients and propose that case-specific alterations can be exploited for classification purposes. + + Introduction Cancer is a complex disease with considerable heterogeneity, both between as well as within cancer types. Finding robust and easily accessible biomarkers is crucial to improve patient well-being and outcome. Such biomarkers can either assist in early detection of cancer, enabling treatment to start at an earlier stage, or they can support diagnosis, treatment selection or response monitoring. As an alternative for biomarkers that require invasive tissue biopsies, liquid biopsy-based biomarkers such as circulating nucleic acids have emerged as a promising tool for cancer diagnosis and monitoring. To date, most studies on circulating nucleic acids in cancer patients have focused on circulating tumor DNA (ctDNA). The amount of tumor DNA in circulation is highly dependent on the tumor type and burden 1 , suggesting that ctDNA can be less informative for certain cancer types or early-stage cancers. Cell-free RNA (cfRNA) may complement ctDNA by reflecting dynamic changes in gene expression during cancer development and progression, or upon treatment, in both healthy and diseased cells. A recent study showed that different cell types contribute RNA to the healthy cell-free plasma transcriptome, and that cfRNA allows monitoring of cell-type-specific changes in diseases, such as non-alcoholic fatty liver and Alzheimer's disease 2 . The potential of cfRNA as prognostic biomarker has also been demonstrated for pre-eclampsia and organ damage in pregnancy 3 . Some studies have also shown potential of using cfRNA biomarkers in certain cancer types 4-6 . Larson et al. 4 specifically looked at genes that are "dark" (no or few detected transcripts) in plasma from healthy controls but abundant in plasma from lung and/or breast cancer patients and found indications of tissue-of-origin signal. Chen et al. 5 reported potential of combining human and microbe-derived plasma cfRNA for distinguishing healthy donor and cancer plasma samples (colorectum, stomach, liver, lung, and esophageal cancer) based on profiles of RNAs showing group-level differences. Roskams-Hieter et al. 6 showed plasma cfRNA classification potential for liver cancer and multiple myeloma patient samples compared to non-cancer controls, with a higher accuracy using learning vector quantization feature selection compared to group-level differential abundance. The authors, however, indicated that liver and bone marrow are major contributors to the plasma transcriptome which may result in more cancer signal. Here, we further examined the impact of different cancer types on plasma cfRNA profiles in different cohorts (Table 1). We first quantified cell-free mRNAs in different cohorts of blood plasma from cancer patients with locally advanced or metastatic tumors and cancer-free donors to uncover tumor or tissue-of-origin mRNA signatures with diagnostic potential. Results revealed substantial heterogeneity in cell-free mRNA abundance among cohorts, questioning the biomarker potential of cell-free mRNA signatures. However, patient-specific alterations, reflected by the number of cell-free mRNAs with differential abundance in a patient sample compared to a cancer-free control population, enabled to accurately distinguish cancer patients from healthy individuals. Table 1: Cohort overview. Name, number of donors (one sample per donor), biofluid type, library preparation method and application of the cohort in this study. (*) eight control donors also included in pan-cancer cohort, data of these donors not used for validation analyses (see Methods). cohort name cancer patients (cancer types) controls biofluid library preparation application pan-cancer cohort 200 (25) 8 plasma mRNA capture (Illumina DNA Prep with Enrichment) discovery three-cancer cohort 36 (3) 30* plasma mRNA capture (Illumina DNA Prep with Enrichment) validation pan-cancer analyses & discovery tail gene concept lymphoma cohort 43 (2) 22 plasma total RNA (SMARTer Stranded Total RNA-Seq Kit v3 -Pico Input Mammalian) validation tail gene concept bladder cancer cohort 12 (1) 12 urine mRNA capture (TruSeq RNA Exome) validation tail gene concept Results A diverse repertoire of cell-free mRNAs in plasma To characterize the blood plasma cell-free mRNA transcriptome, we collected blood plasma from a pan-cancer discovery cohort consisting of 208 individuals covering 25 different cancer types and aged-matched healthy controls (Figure 1a, Supplemental Table 1). MRNA capture sequencing of the pan-cancer plasma cohort resulted in 3 to 56M paired-end reads per sample (median 16M) after quality filtering with no significant differences among groups (Kruskal-Wallis test p>0.05) (Supplemental Figure 1). On average 84% (95% CI [82.59, 84.66]) of uniquely mapped reads were assigned to protein coding genes and most other reads originated from mitochondrially encoded ribosomal RNAs (Supplemental Figure 2). Per sample, between 7439 and 13,039 unique mRNAs (median 10,932) were detected with at least 10 counts -we use the term mRNA to refer to all mRNA reads mapping to the same gene (Figure 1b). All plasma samples had 5263 mRNAs in common and 9908 mRNAs were found in at least 75% of the samples. Of note, we detected more mRNAs in plasma from acute myeloid leukemia (AML) patients compared to plasma from controls and from patients with solid cancers (Kruskal-Wallis p=6.6E-4, effect size= 0.163, post hoc two-sided Wilcoxon rank-sum q < 0.05 for AML vs controls and 16 out of 24 solid cancer types). Although some AML samples were sequenced deeper, sequencing depth alone did not explain the higher number of detected mRNAs (Supplemental Figure 3). We next determined the mRNA concentration using synthetic RNA spikes that were added to each lysed plasma sample (Figure 1c). MRNA concentrations ranged from 0.003 to 8.302 ng/ml with an average concentration of 0.486 ng/ml. Despite the higher number of detected mRNAs, the mRNA-concentration in AML plasma was not significantly higher than controls or other cancer types. Moreover, though the largest median mRNA concentration (READ, 0.648 ng/ml) was more than 10 times higher than the smallest median (ANA, 0.063 ng/ml), no significant concentration differences were found between the groups (Kruskal-Wallis test p=0.0275, effect size=0.087, with all post hoc two-sided Wilcoxon rank-sum q > 0.05). Besides the small sample sizes, this may be related to the considerable cfRNA concentration variability between patients, which is also reported at cfDNA level 7 . We then searched for evidence of tumor-derived cfRNA by investigating known cancer fusion transcripts in individual plasma samples. In every cancer plasma sample, we only considered fusion transcripts that had been documented in tumor tissue of that same cancer type. Such fusion transcripts were only identified in plasma samples of AML patients. The PML::RARA fusion transcript 8 was found in two AML plasma samples with several reads supporting the fusion (7 to 19 spanning pairs and 6 to 9 unique spanning reads). The reciprocal RARA::PML fusion was found in one of both AML plasma samples (19 pairs and 17 unique reads). In plasma samples of solid cancer types, we only detected fusion transcripts that had not been reported in the tumor tissue of these cancer types (Supplemental Table 2). Of note, these fusion transcripts had fewer supporting reads (1 to 2 pairs and 2 to 5 unique reads) than the PML::RARA/RARA::PML fusions in AML and may thus represent false positives. Moreover, other fusions with few supporting reads were also identified in healthy control plasma samples. While the two AML samples with the PML::RARA fusion transcripts were sequenced deeper than other samples, the fusions were still detected when downsampling to the median number of reads of the pan-cancer cohort. These analyses confirm the presence of tumor cell-derived cfRNA in plasma samples from AML patients. Figure 1: Pan-cancer plasma cohort overview, mRNA content and sample clustering. a, Primary sites of the 25 cancer types in the pan-cancer cohort. Primary sites are indicated with colored dots and corresponding cancer types are listed (top to bottom) with the (sub)type abbreviations between brackets. Full names and more details can be found in Supplemental Table 1. b, Number of plasma mRNAs, detected with at least 10 counts, across cancer types and healthy controls. Groups ordered according to median. c, Variability in plasma mRNA concentration across cancer types and healthy controls. Groups ordered according to median. b and c, Boxplots show lower quartile (Q1), median, and upper quartile (Q3). Whiskers extend from the lower and upper quartile to the smallest and largest value, respectively, within at most 1.5*interquartile range (Q3-Q1) from that quartile. More extreme points are plotted as individual dots. d, Principal component analysis using the 500 most variable mRNAs in the pan-cancer cohort (with variance stabilizing transformation DESeq2). Yellow squares: control samples; red triangles: acute myeloid leukemia samples; blue dots: non-leukemia cancer samples. e and f, Principal component analysis using the 500 most variable mRNAs in controls and anaplastic astrocytoma (ANA) or prostate cancer (PRAD) patients, respectively. Replicate numbers are indicated in the plot. PC: principal component. Differential mRNA abundance in cancer patients' plasma compared to controls Principal component analysis revealed a clear separation between AML and other plasma samples for six out of eight AML patients (Figure 1d). Control samples mostly clustered in between samples of solid tumors. Analysis of individual solid tumor types revealed that most patient samples could be separated from controls in the first two principal components (Figure 1e and f, Supplemental Figure 4), suggesting that plasma cfRNA profiles of cancer patients differ from that of controls. Differentially abundant mRNAs were identified in every cancer-control comparison, with numbers varying from 10 to 2583 (median 128) for higher abundant mRNAs and from 89 to 2727 (median 221) for lower abundant mRNAs in cancer compared to control (Figure 2a). Most differentially abundant mRNAs were found in AML (5310 mRNAs), followed by uterine cancer (UCEC) and sarcoma (SARC) with 1154 and 763 mRNAs, respectively. Head and neck (HNSC), rectal (READ) and breast cancer (BRCA) had the lowest number of differentially abundant mRNAs (127, 136, and 145 respectively). In total, 6758 unique mRNAs were differentially abundant in one or more cancer-control comparisons (b) and the median absolute fold change was 3.1. One quarter of differentially abundant mRNAs had an absolute fold change of at least 4.6. Hierarchical clustering based on differential mRNA abundance mostly separated cancer and control samples but also revealed variability in gene abundance within groups (Figure 2b and c, Supplemental Figure 5). 4442 mRNAs (66%) were differentially abundant in one cancer type only (Figure 2d), and most (3680 mRNAs) were identified in AML. While many mRNAs were differentially abundant in plasma of only one or a few cancer types, 26 differentially abundant mRNAs were recurrent in 23 out of 25 cancer-control comparisons (Figure 2d). These mRNAs were all lower abundant in cancer compared to controls and showed enrichment for inflammatory and immune response pathways (Supplemental Table 3). Figure 2: Differentially abundant mRNAs in cancer versus control. a, Differentially abundant mRNAs in plasma from cancer patients versus controls. Cancer types ordered based on total number of differentially abundant mRNAs (q < 0.05 and |log2 fold change|>1). b and c, Volcano plot and heatmap of differentially abundant mRNAs in anaplastic astrocytoma vs controls (b) and in prostate cancer vs controls (c). Volcano plots have a horizontal line at q = 0.05 and vertical lines at log2 fold change of -1 and 1, resp. Higher (red): higher abundant in cancer compared to control samples (q < 0.05 and log2 fold change > 1); lower (blue): lower abundant in cancer compared to control samples (q < 0.05 and log2 fold change < -1). Heatmaps show relative abundance of differentially abundant mRNAs (columns) in individual plasma samples (rows) based on log2(normalized counts+1) followed by z-score transformation per gene. Clustering of genes and samples based on Pearson correlation. d, Recurrence of differentially abundant mRNAs (shown in a) across cancer types. Enrichment of pathways and cell types in cancer patients' plasma To explore the pathways, cell types and tissues underlying the differentially abundant mRNAs in cancer plasma, we first applied gene set enrichment analyses using hallmark 9 , Kyoto Encyclopedia of Genes and Genomes (KEGG) 10 , and custom tissue-and cell type gene sets (Figure 3a, Supplemental Figure 6a and b). We observed a negative enrichment of various immune-related signals (interleukin signaling, interferon gamma and inflammatory response, and neutrophils) across all cancer types except UCEC, with q<0.05 in at least 21 out of 25 cancer types. 'Ribosome' was also negatively enriched in all solid cancer types (q<0.05 in 19 out of 24 types). These results are in line with a previous study by Chen et al. 5 on plasma cfRNA profiles of five early-stage solid cancer types. Other negatively enriched gene sets with q<0.05 in at least half of the cancer types in our cohort were 'epithelium' and 'blood' related. Conversely, 'extracellular matrix receptor' and 'focal adhesion' gene sets were significantly positively enriched in at least half of the cancer types, confirming results by Chen et al. Other positively enriched gene sets included 'epithelial-mesenchymal transition', 'vasculature', 'adipose tissue', 'brain', and 'hematopoietic stem cells'. Cancer type-specific enrichment of cfRNA reflecting the primary tumor site was only observed for liver cancer patients. 'Liver tissue', 'liver cancer tissue', and 'hepatocyte' gene sets were all positively enriched in liver cancer patients' plasma (q<0.05, Figure 3b) and not in plasma from any other cancer type. A 'bronchial epithelial cells' gene set was significantly enriched in lung adenocarcinoma and lung squamous cell carcinoma (LUAD q=0.027; LUSC q=0.026), but this enrichment was also found in four other cancer types (ANA, CESC, CHOL, and UCEC). We then applied computational deconvolution to enumerate the cell type fractions contributing to plasma cfRNA profiles (Supplemental Table 4). Platelets, erythrocytes/erythrocyte progenitors, monocytes, macrophages, and B-cells formed the top-5 of most abundant cell types that contributed to plasma cfRNA of both controls and patients with solid tumors (Figure 3c). Except for macrophages, these cell types were also reported as the most abundant cell types in a previous study on healthy donor plasma 2 . Plasma from AML patients showed a very different composition. Several cell types -including hematopoietic stem cells, myeloid progenitor cells, and monocytes -were increased or even exclusively present in AML plasma samples, in line with the blood cell counts typically observed in these patients 11 . Myeloid progenitor cells, normally not present in peripheral blood, were the second most abundant cell type in the AML plasma samples (mean 10.7%), with almost no signal in other plasma samples (mean 0.1%) (Supplemental Figure 7). Moreover, the platelet fraction in AML plasma was significantly lower compared to other groups (two-sided Wilcoxon rank sum p=3.8E-5, effect size=0.289) with on average 18.3% in AML compared to 62.3% in other samples, reflecting the reduced thrombocyte counts commonly found in leukemia patients 11 . In plasma from patients with solid tumors, the platelet fraction was significantly higher compared to controls (Supplemental Figure 7, two-sided Wilcoxon rank-sum p=2.3E-2, effect size=0.163), which is in line with the prognostic value of platelet counts and the frequent occurrence of thrombocytosis in solid tumor patients 12,13 . For neutrophils, erythrocyte/erythroid progenitor cells, thymocytes, and T-cells, the fraction was significantly lower in cancer patients' plasma compared to controls (two-sided Wilcoxon rank-sum p=1.8E-2 (effect size=0.169), p=1.4E-2 (effect size=0.176), p=1.8E-2 (effect size=0.164), p=4.3E-4 (effect size=0.252), respectively). Figure 3: Gene set enrichment and cell type deconvolution reveal pan-cancer and cancer-type specific signals. a, Gene set enrichment analysis results for custom tissue and cell type sets. Only sets that are significantly enriched in at least half of the cancer types are shown. Gene sets with q < 0.05 colored according to normalized enrichment scores (NES). b, Liver-related gene set enrichment in plasma from liver cancer patients, gene sets with q < 0.25 colored according to NES. Detailed information about gene sets can be found in Methods. c, Average fraction of the five most abundant cell types based on deconvolution. Other: cumulative sum of other cell types. The five most abundant cell types were selected based on mean ranking in all groups. Independent validation cohort confirms systemic signal and cfRNA heterogeneity To validate our findings, we processed an independent (three-cancer) cohort of plasma samples which, after removing samples with low read depth, included samples from 20 cancer-free control donors and 35 cancer patients with one of three different solid cancer types: prostate (PRAD, n=12), ovarian (OV, n=11), and uterine (UCEC, n=12) cancer. 7 to 61M paired-end reads were obtained per sample (median 28M) with no significant difference in sequencing depth among groups (Kruskal-Wallis p=0.7858) (Supplemental Figure 8a). In line with the pan-cancer cohort, we detected between 8642 and 11,944 mRNAs per sample (Supplemental Figure 8b), with 6194 mRNAs shared across all plasma samples of this three-cancer cohort. MRNA concentrations ranged from 0.007 to 0.414 ng/ml (Supplemental Figure 8c). Most cancer samples could be separated from controls in the first principal component, especially for prostate and ovarian cancer (Figure 4a). Based on differential abundance analyses between cancer and control samples, we found the highest number of differentially abundant mRNAs in OV (995 higher abundant, 1644 lower abundant), followed by PRAD (848 higher abundant, 1043 lower abundant) (Figure 4b). The lowest number of differentially abundant mRNAs was found for UCEC (243 higher abundant, 349 lower abundant) even though this cancer type had the most differentially abundant mRNAs in the pan-cancer cohort. Hierarchical clustering based on differential mRNA abundance in this three-cancer cohort mostly separated cancer and control samples but again revealed variability in mRNA abundance within groups (Supplemental Figure 9). 189 and 299 mRNAs were significantly higher and lower abundant, respectively, in plasma from all three cancer types compared to controls (Figure 4b). The lower abundant mRNAs included all 26 mRNAs that were identified as lower abundant in most cancers in the pan-cancer cohort. There was a significant, although limited, overlap in differentially abundant mRNAs between the three-cancer and the pan-cancer cohort for each of the three cancer types (Fisher's exact test on lower abundant mRNAs: OV p=8.4E-118 (odds ratio=30.0), PRAD p=1.1E-155 (odds ratio=59.3), UCEC p=7.6E-13 (odds ratio=4.2); on higher abundant mRNAs: OV p=4.0E-4 (odds ratio=4.6), PRAD p=2.9E-52 (odds ratio=12.2), UCEC p=2.2E-5 (odds ratio=3.3); Jaccard index [0.9-14.3%]) (Figure 4c and Supplemental Figure 10a). Respectively 38%, 49%, and 95% of the differentially abundant mRNAs in OV, PRAD, and UCEC in the pan-cancer cohort could not be confirmed in the three-cancer cohort. Notably, mRNAs with lower abundance showed a higher overlap between both cohorts than mRNAs with higher abundance. In line with the pan-cancer cohort, there was no enrichment of cancer type specific gene sets reflecting tumor tissue of origin in the OV, PRAD and UCEC samples in the three-cancer cohort. However, enriched gene sets shared among at least half of the cancer types in the pan-cancer cohort were confirmed in the three-cancer cohort, except for adipose tissue (Figure 4d, Supplemental Figure 10b and c). Cell type deconvolution confirmed platelets, erythrocyte/erythroid progenitor cells, monocytes and B-cells among the top-5 cell types contributing cfRNA to plasma (Figure 4e, Supplemental Table 4), with neutrophils being ranked as the fourth most abundant cell type. The higher platelet fraction in cancer patient plasma was even more pronounced in the three-cancer cohort (two-sided Wilcoxon rank-sum cancer versus control p=5.8E-9, effect size= 0.679), especially for ovarian and prostate cancer (Supplemental Figure 11). Neutrophils, erythrocyte/erythroid progenitor cells, thymocytes and T-cells again had a lower plasma fraction in cancer patients compared to controls (two-sided Wilcoxon rank-sum p=4.4E-6 (effect size=0.556), p=3.2E-7 (effect size=0.610), p=6.4E-5 (effect size=0.419), p=4.1E-2 (effect size=0.261), respectively). Taken together, the limited overlap of differentially abundant mRNAs between the pan-cancer and three-cancer cohort highlights plasma transcriptome variability, yet enrichment and cell type contribution analyses confirm the systemic cfRNA changes in plasma from cancer patients. Figure 4: Three-cancer cohort confirms findings from pan-cancer cohort and reveals significant, yet limited, overlap in differentially abundant mRNAs. a, Principal component analysis on top 500 most variable mRNAs shows distinct clustering of most cancer and control samples (variance stabilizing transformation DESeq2). b, 0verlap between differentially abundant mRNAs detected in individual cancer-control comparisons. Circles are proportional to the number of genes. c, Overlap between differentially abundant genes in prostate cancer versus control for the pan-cancer and three-cancer cohort. Fisher's exact test: p=2.9E-52, odds ratio=12.2, Jaccard index=8.3% for higher abundant mRNAs in cancer versus control (higher); p=1.1E-155, odds ratio=59.3, Jaccard index=14.3% for lower abundant mRNAs (lower). Circles are proportional to the number of genes. d, Gene set enrichment analysis results for custom tissue and cell type sets. Only pathways of Figure 3a shown. Gene sets with q < 0.05 colored according to normalized enrichment scores (NES). e, Average fraction of the five most abundant cell types and macrophages based on cell type deconvolution. Other: cumulative sum of other cell types. Differential abundance: q<0.05 & |log2 fold change|>1. OV: ovarian cancer; PRAD: prostate cancer; UCEC: uterine cancer. cfRNA tail genes distinguish cancer samples from controls Apart from liver cancer and AML, our analyses did not reveal evidence for a tumor or tissue of origin signal among mRNAs that were more abundant in cancer patient versus control plasma. Moreover, higher abundant mRNAs were less recurrent across cancer types and showed lower validation rates between cohorts compared to mRNAs with lower abundance in cancer versus control plasma. Our analyses suggest that the latter represent a systemic immune response rather than a tumor derived signal. Based on these observations, we hypothesized that (tumor derived) cfRNA signals may be heterogeneous among patients, and that classic differential mRNA and pathway abundance analysis between groups may not reveal such non-uniform signals. We therefore decided to investigate cfRNA profiles in individual cancer patients using samples from the three-cancer cohort. To capture aberrations in the cfRNA profile of an individual cancer patient, we compared the mRNA abundance of a cancer plasma sample to the mRNA abundance distribution in the entire control group, that served as a reference. More specifically, a z-score was calculated for each mRNA based on the mean and standard deviation of the log2 normalized count distribution in the control group. Abundant mRNAs with an absolute z-score of at least 3 were defined as "tail genes" for that patient, referring to their position in one of the tails of the control z-score distribution (Figure 5a). We searched for tail genes in each cancer patient from the three-cancer cohort (where we could take advantage of a large control group) and in each control sample, always using all other control samples as reference (Supplemental Figure 12). We identified a total of 3312 unique tail genes in this cohort and observed a significantly higher number of tail genes in plasma samples from cancer patients compared to controls (Kruskal-Wallis p=0.0012, effect size=0.222, Figure 5b). 74% of the tail genes were not identified as differentially abundant in cancer samples versus controls (Figure 5c). Moreover, for 40.0% of cancer sample tail genes that were differentially abundant between cancer and controls, the direction (i.e. positive or negative z-score) in at least one cancer sample was opposite to the direction reported by the differential abundance analysis, further highlighting the heterogeneity among patients. Notably, the recurrence of tail genes was much higher in cancer samples than controls; 98% of the control tail genes were identified in only one control sample while 47% of cancer tail genes were identified in more than one cancer sample (Figure 5d). To determine the significance of the observed results, we randomly swapped sample labels prior to tail gene identification and repeated this procedure 20 times. Doing so, we did not observe a significant difference in number of tail genes between the newly labeled cancer samples and controls in any of the repeats (all Kruskal-Wallis p≥0.05), confirming that an increased number of tail genes is specifically associated with cancer samples. Tail genes detected in one cancer type showed significant overlap with tail genes detected in another cancer type (Fisher's exact tests p≤5.4E-168, odds ratio: [11.1-15.2]), yet limited similarity (Jaccard index: [19.7-34.8%]) (Figure 5e). The 1182 unique control tail genes also showed significant overlap with any of the cancer tail gene subsets (pairwise Fisher exact tests p: [5.9E-15 -8.5E-32], odds ratio: [2.1-2.5]) with less similarity (Jaccard index: [7.4-11.9%]). 63% of control tail genes were not identified as tail gene in any of the cancer samples. Seven Cancer Gene Census oncogenes were identified as tail genes (z>3) in at least one sample, which is significantly more than expected by chance (Fisher's exact p=0.019, odds ratio=3.13). While most oncogenes were only identified as tail gene in a single sample, KAT6A was found in seven cancer samples (4 OV, 2 PRAD, and 1 UCEC) but also in one control sample. Overrepresentation analyses (Supplemental Figure 13, Supplemental Table 5) demonstrated that tail genes that are higher abundant in at least one cancer sample (z>3) were involved in different processes including platelet activation and signaling, actin binding and dynamics, and extracellular matrix. Tail genes that are lower abundant in at least one cancer sample (z<-3) were mainly related to mRNA splicing and translation. Significantly enriched processes in the cancer tail gene sets were not enriched in the control tail gene set, and processes enriched in the lower abundant (cancer) tail genes showed higher recurrence across the three cancer types compared to those enriched in the higher abundant tail genes. Given the higher number of tail genes in cancer versus controls, we further assessed the binary (cancer/control) classification potential of the number of tail genes per sample. We first selected a subset of tail genes, further coined biomarker tail genes, that were significantly associated to either the cancer or control state using leave-one-out Fisher exact tests (see Methods for details). This resulted in 108 biomarker tail genes, with a significant difference in number of biomarker tail genes in cancer and control samples (two-sided Wilcoxon rank-sum p=3.1E-11, effect size=0.845). None of the biomarker tail genes were recurrent tail genes in control samples and 54% were not identified as differentially abundant in any cancer type compared to controls (Supplemental Figure 14). Applying the biomarker tail gene selection strategy to the 20 datasets with randomly swapped sample labels resulted in considerably fewer or no biomarker tail genes at all: between 0 and 7 biomarker tail genes (median: 2) instead of 108. When selecting biomarker tail genes in a specific cancer group versus controls, we obtained 143 biomarker tail genes for OV, 46 for PRAD, and 13 for UCEC. The difference in number of biomarker tail genes between cancer and control samples per biomarker tail gene subset was more significant than based on all tail genes (two-sided Wilcoxon rank-sum: for OV p=1.1E-7 and effect size=0.864; for PRAD p=5.7E-8 and effect size=0.872; for UCEC p=1.4E-8 and effect size=0.912) (Figure 5f). Note that the cancer type-specific biomarker tail gene sets showed significant overlap (Fisher's exact p≤1.8E-3, odds ratio: [10.1-22.6]) with limited similarity (Jaccard index: [2.6-9.9%]) (Figure 5h). We then built a classifier for discriminating cancer and control samples based on the number of biomarker tail genes in a particular sample. Using leave-one-out cross-validation, we obtained perfect binary classification for ovarian or prostate cancer samples and controls (AUC=1.000, based on 143 and 46 biomarker tail genes, resp.), and a high classification performance for uterine cancer samples and controls (AUC=0.917, based on 13 biomarker tail genes) (Figure 5g). The classification performance was still high when combining all cancer samples (AUC=0.980, based on 108 biomarker tail genes). Together, these analyses suggest that individual cancer samples can be distinguished from controls based on the number of biomarker tail genes in their plasma cfRNA profiles. Figure 5: Higher number of tail (biomarker) genes in cancer patient plasma samples compared to controls. a, Tail gene identification workflow. A gene in a particular sample for which the abundance deviates more than 3 standard deviations from the reference group abundance distribution is a tail gene for that sample. The number of tail genes is used as a metric to determine cancer/control status. b, Boxplot of the number of tail genes per group, with grey dots representing individual sample counts. Kruskal-Wallis p = 0.001, effect size = 0.222. Two-sided Wilcoxon rank-sum q-values for specific cancer versus control comparisons indicated in the plot. c, Z-score distribution of tail genes in the different samples, all 3312 genes that fulfilled tail gene criteria in at least one of the samples are shown. For visualization purposes, z-scores above 5 or below -5 were reduced to 5 and -5, respectively. Genes (rows) clustered based on Euclidean distance. d, Number of tail genes shared by a certain fraction of samples of the same type. Number of samples indicated between brackets. e, Overlap between tail genes detected in the different cancer types. f, Boxplot of number of biomarker tail genes per subset in control (CONTROL) versus the respective cancer samples. Total number of biomarker tail genes considered for each comparison: 108 for all cancer (n=35) versus control, 143 for OV versus control, 46 for PRAD versus control, 13 for UCEC versus control. Grey dots represent individual sample counts. Two-sided Wilcoxon rank-sum p-values indicated in plot. g, ROC curve of leave-one-out cross-validated classifier for binary cancer/control classification based on the number of biomarker tail genes for control and ovarian cancer samples (OV), prostate cancer samples (PRAD), uterine cancer samples (UCEC), or all cancer samples (all) using the respective biomarker tail gene sets. h, Overlap of cancer type-specific biomarker tail genes. Boxplots show lower quartile (Q1), median, and upper quartile (Q3). Whiskers extend from the lower and upper quartile to the smallest and largest value, respectively, within at most 1.5*interquartile range (Q3-Q1) from that quartile. More extreme points are plotted as individual black dots. 62 plasma samples considered in total from 27 cancer-free controls (CONTROL), 11 ovarian cancer (OV), 12 prostate cancer (PRAD), and 12 uterine cancer patients (UCEC). To independently validate the classification potential of the number of biomarker tail genes, we generated additional plasma cfRNA profiles (lymphoma cohort), including plasma samples from 22 cancer-free control donors and 43 lymphoma patients, more specifically 30 diffuse large B-cell lymphoma (DLBCL) and 13 primary mediastinal large B-cell lymphoma (PMBCL) patients. We identified 5374 unique tail genes, again with more tail genes in cancer samples compared to controls (Kruskal-Wallis p=2.9E-5, effect size= 0.305, Supplemental Figure 15a). All control tail genes were identified in only one control sample while 45.9% of DLBCL and 44.2% of PMBCL tail genes were identified in more than one DLBCL and PMBCL sample, respectively (Supplemental Figure 15b). When swapping sample labels (20 repeats) prior to tail gene identification, no significant differences in the number of tail genes between cancer and control samples were observed in any of the repeats (Kruskal-Wallis p≥0.05). Among the tail genes, 297 and 931 were identified as biomarker tail genes using leave-one-out Fisher's exact tests for respectively DLBCL or PMBCL. The number of biomarker tail genes was significantly higher in DLBCL and PMBCL compared to control samples (two-sided Wilcoxon rank-sum: for DLBCL p=3.3E-9, effect size=0.822; for PMBCL p=1.7E-6, effect size=0.811, Figure 6a). A binary classifier (cancer/control) based on biomarker tail genes with leave-one-out cross-validation resulted in an AUC of 0.937 and 0.944 for DLBCL and PMBCL, respectively (Figure 6b). 78% of the biomarker tail genes were not differentially abundant in any lymphoma subtype versus controls. Notably, early-stage lymphoma patients also displayed an increased number of biomarker tail genes (Figure 6c). Cancer diagnosis (i.e. DLBCL or PMBCL) was the only factor significantly associated to the number of biomarker tail genes (p=1.8E-4 and 3.1E-5 for DLBCL and PMBCL, respectively) in a generalized linear model including diagnosis and tumor stage. To test if our cancer-control classification strategy based on biomarker tail genes is also applicable to other biofluids, we applied it to a cohort of urine cfRNA profiles (bladder cancer cohort) 14 . This cohort included urine samples of 12 bladder cancer patients (BLCA) and 12 cancer-free control donors. We identified 1152 tail genes of which 39 qualified as biomarker tail gene. The number of biomarker tail genes was significantly higher in bladder cancer versus controls (two-sided Wilcoxon rank-sum p=6.0E-5 and effect size=0.826, Figure 6d) and 62% of biomarker tail genes were not differentially abundant between the cancer and control group. In line with the other datasets, sample label swapping prior to tail gene identification did not result in significant differences in the number of tail genes between the newly labeled cancer samples and controls. A binary classifier based on biomarker tail genes with leave-one-out cross-validation resulted in an AUC of 0.903 (Figure 6e). Taken together, our results confirm that the number of biomarker tail genes in blood plasma cfRNA enables accurate classification of cancer patients from controls and suggest that the concept may also be applicable to cfRNA profiles from other biofluids. Figure 6: Number of biomarker tail genes is significantly higher in cancer versus control samples in an independent blood plasma and urine cohort. a-c, lymphoma cohort: plasma samples from 22 control, 30 diffuse large B-cell lymphoma (DLBCL), and 13 primary mediastinal large B-cell lymphoma (PMBCL) patients. a, Boxplot of number of biomarker tail genes in control versus the respective cancer samples. Total number of biomarker tail genes considered for each comparison: 297 for DLBCL versus control, 931 for PMBCL versus control. Grey dots represent individual sample counts. Wilcoxon rank-sum p-values indicated in plot. b, ROC curve of leave-one-out cross-validated classifier for binary cancer/control classification based on the number of biomarker tail genes for control and DLBCL, or control and PMBCL using the respective biomarker tail gene sets. c, Number of biomarker tail genes of panel a visualized per cancer stage. d-e, bladder cancer cohort: urine samples from 12 control (CONTROL) and 12 bladder cancer (BLCA) patients. d, Boxplot of number of biomarker tail genes in control versus cancer samples. 39 biomarker tail genes considered in total. Wilcoxon rank-sum p-value indicated in the plot. e, ROC curve of leave-one-out cross-validated classifier for binary cancer/control classification based on the number of biomarker tail genes. Boxplots: lower quartile, median, upper quartile, whiskers of 1.5*interquartile range; more extreme points are indicated by black dots. Discussion In this study, we explored cancer-induced changes in blood plasma cfRNA profiles and investigated if these changes could be exploited for classification purposes. Results from our pan-cancer discovery cohort revealed that AML was the only cancer type for which we found direct evidence for tumor-derived cfRNA in plasma in the form of PML::RARA fusion transcripts. Also, deconvolution of AML plasma cfRNA revealed a high fraction of myeloid progenitor cells, providing additional -albeit indirect -evidence for tumor-derived cfRNA. For solid tumor types, we identified differentially abundant mRNAs in plasma for every cancer versus control comparison, but a cancer type-specific signal was only observed in plasma from liver cancer patients through the enrichment of liver-specific gene sets. Liver is a major contributor to the plasma transcriptome 4 and its high vascularization and blood flow may facilitate mRNA release in blood upon liver cancer progression. Increased cfDNA contributions from the liver have also been observed in liver cancer patients 15 . Note that, despite the enrichment of liver-related mRNAs, the total cfRNA concentration was not increased in plasma from liver cancer patients, indicating that this signal was still strongly diluted. Considering the organ contribution and dilution argument, other types of liquid biopsies may be better suited to monitor tumor-derived cfRNA for other cancer types. For instance, we could not detect enrichment of prostate tissue-related nor prostate cancer-specific transcripts in blood plasma from prostate cancer patients, in contrast to earlier findings in seminal plasma samples 16 . We did observe several general trends in patients' blood plasma transcriptomes in the pan-cancer cohort, which were confirmed in an independent cohort. The lower abundance of immune-related mRNAs may be linked to the suppressed immune system status in cancer patients. It was recently shown that local tumors can have a systemic impact on the immune system, and that different tumor sites of origin resulted in distinct relative abundances and activity profiles of immune cell types 17 . This may be linked to the observed variations in immune-related mRNA abundance, both between and within cancer types. The (immune) cell-related mRNA contributions agreed with reported cell abundance trends found in blood from several cancer patients 18 , with the exception of neutrophil-related mRNAs that are lower abundant in plasma. However, neutrophils can have a dual role in cancer 19 and reduced neutrophil cell-free mRNA abundance may result from neutrophils specifically recruited to tumor sites. Finally, the negative enrichment of blood/erythrocyte gene sets may be linked to anemia, a common condition in cancer patients. Among the processes that we found positively enriched in cancer patient plasma, some are linked to hallmarks for cancer progression and metastasis, including epithelial-to-mesenchymal transition and increased blood vessel signal that stem from leaky vasculature and angiogenesis 20 . A limitation of the pan-cancer cohort is that control samples were not always perfectly age and sex-matched. The anaplastic astrocytoma, cervical cancer, and testicular germ cell tumor patients were younger as these types predominantly appear at younger age. A dedicated cohort with matched control samples may be better for exposing cancer-related changes in these cancer types. Similarly, while male and female donors were balanced in the control group, this was not always possible for the different cancer types due to sex-specificity (e.g. prostate, testicular, breast, cervical, and ovarian cancer) or sample availability. Nevertheless, at most 4 out of 8 donors were not sex-matched in cancer vs control group comparisons in the pan-cancer cohort, and our main conclusions were not impacted when only using controls of the same sex in the three-cancer cohort. Another limitation is the relatively small sample size per cancer type, which might contribute to false positives and insufficient power to detect true differences. The variable mRNA abundance in plasma, combined with the limited overlap of differentially abundant mRNAs across cohorts of the same cancer type prompted an alternative approach that can effectively deal with patient heterogeneity. The tail gene concept satisfies this requirement by identifying mRNAs whose abundance in individual samples significantly deviates from the abundance distribution in a reference group. While the dark channel biomarker gene concept 4 is in part based on a more patient-centric approach, it also requires differential abundance at group level to be considered potential biomarkers. Moreover, the dark channel concept specifically looks at genes that are absent (dark) in control samples and therefore cannot capture changes in transcript abundance of genes that are also present in healthy plasma transcriptomes. Of note, 19 dark channel biomarker genes were identified for lung and breast cancer by Larson et al. 4 but five of those genes could be readily detected in control samples from our cohorts, indicating that the "darkness" (i.e. absence of gene transcripts in control samples) should be re-evaluated in different cohorts. Tail gene counting is a simple metric for classifying samples as cancer or controls. Considering that it is not based on gene signatures with weighted contributions of individual features, we believe it could be more robust and compatible with observed patient heterogeneity. Nevertheless, it requires a large and representative group of cancer-free control samples. A control group only needs to be established once, if the entire workflow -from sample collection to RNA-sequencing -is standardized across cancer and control samples. This approach should minimize the introduction of batch effects that may skew cfRNA distributions between a pre-established cohort of cancer-free controls and newly profiled cancer or control samples. In this regard, our group has demonstrated that preanalytical variables (blood collection tube, time between blood collection and plasma preparation, RNA extraction kit) can indeed dramatically impact cfRNA profiles 21 . Deviations in the number of tail genes is not necessarily restricted to cancer patients, nor to blood plasma. Indeed, cfRNA from urine provides a good source of biomarker tail genes to identify bladder cancer patients. Other diseases may be characterized by an increase in tail genes, although the specific tail (biomarker) genes may differ. Further studies are needed to examine the applicability and overlap of tail genes in non-malignant disease cohorts. Similarly, while we have preliminary evidence that biomarker tail genes are also significantly more numerous in low stage patients, additional follow-up studies comprising different stages across various cancer types could reveal whether the concept is suited for early-stage cancer screening or monitoring. Methods Sample cohorts The study was approved by the ethics committee of Ghent University Hospital, Belgium (no. B670201734362) and performed in accordance with the Declaration of Helsinki. Pan-cancer plasma and three-cancer cohort Blood plasma samples of 266 human donors were obtained from Proteogenex (CA, USA) with informed consent from all donors. The pan-cancer discovery cohort covers 25 different cancer types and a cancer-free control group. For each type, plasma was collected from 8 different donors. The three-cancer cohort consisted of prostate cancer (PRAD), ovarian cancer (OV), and endometrial cancer (UCEC) plasma samples, with 12 donors per type. This cohort also included plasma samples from the 8 control donors of the pan-cancer cohort as well as from 22 additional control donors. For independent validation of pan-cancer results, only the control samples from the additional control donors were used. For tail gene discovery, all control samples in the three-cancer cohort were used. All cancer patients in both cohorts had locally advanced (stage 3) to metastatic cancer (stage 4), except for 6 seminoma patients (stage 1). Blood was drawn before treatment with 6 exceptions: 4 glioblastoma patients (grade 4-5) already received chemo and radiation therapy, one prostate cancer patient with metastasis received hormone therapy, and one melanoma patient with metastasis received immunotherapy. Demographic and clinical information can be found in Supplemental Table 1. Lymphoma cohort The lymphoma cohort consisted of plasma samples from 65 human donors: 30 DLBCL patients, 13 PMBCL patients, and 22 cancer-free control donors. Plasma samples were collected following approval by the ethics committee of Ghent University Hospital, Belgium (B670201733701), and written, informed consent was obtained from all patients. Bladder cancer cohort The bladder cancer cohort is described in more detail elsewhere 14 . This cohort consists of urine samples from 12 muscle-invasive bladder cancer patients and 12 control donors. Blood collection and plasma preparation Blood from the pan-cancer and three-cancer cohort donors was collected in EDTA vacutainer tubes (Becton Dickinson, 367525). Blood was stored at 4 °C until plasma preparation which started by gently inverting the EDTA tube 10 times and centrifugating for 10 min at 1500 g (without brake at 4 °C). Supernatans was then transferred to 15 mL centrifuge tubes and centrifuged again for 10 min at 1500 g (without brake at 4 °C). The resulting platelet depleted plasma was transferred to 2 mL cryovials, frozen and stored at -80 °C within 4 hours of blood collection. Plasma samples were shipped on dry ice and only thawed on ice immediately before RNA isolation. For the lymphoma cohort, 2.5 mL of blood was collected from each donor in PAXgene Blood DNA Tubes (BD Biosciences, 761165). Plasma was prepared by a one-step 15 min centrifugation at 1900 g without brake (room temperature) and stored at -80 °C within 4 hours of blood collection. RNA isolation and gDNA removal RNA was isolated from 200 µL of plasma with the miRNeasy Serum/Plasma kit (Qiagen, 217184), according to the manufacturer's instructions. For the pan-cancer and three-cancer cohort, 2 µL Sequin spike-in controls (1/1,300,000 of stock solutions mix A, Garvan Institute of Medical Research, Australia) was added to the lysate during RNA isolation as previously described 22 . For the lymphoma cohort, 2 µL of Sequin spike-in controls were added to the sample lysates (1/5000 of stock solution mix A (Garvan Institute of Medical Research) and 2 µL of External RNA Control Consortium (ERCC) spike-in controls (ThermoFisher Scientific, 4456740) were added to 12 µl RNA eluate. Genomic DNA was removed by adding 1 μL HL-dsDNase (ArcticZymes, 70800-202) and 1.6 µL reaction buffer (ArcticZymes, 66001) to 12 µL RNA eluate, 10 min incubation at 37 °C, followed by 5 min incubation at 55 °C. RNA was stored at -80 °C and thawed on ice immediately before library preparation. Messenger RNA capture library preparation, sequencing, and quantification MRNA capture library preparation in pan-cancer and three-cancer cohorts started from 8.5 µL DNase treated RNA eluate. cDNA synthesis was performed using TruSeq RNA Library Prep for Enrichment (Illumina, 20020189) as previously described 22 . Briefly, RNA was fragmented, and first strand cDNA was generated using random priming. RNA templates were subsequently removed and replaced by a newly synthesized second strand of cDNA. AMPure XP beads (Beckman Coulter Life Sciences, A63881) were used for purifying the blunt-ended double stranded cDNA. 30 μL cDNA of each sample was then used as input for Illumina DNA Prep with Enrichment (previously Nextera Flex for Enrichment; Illumina, 20025524). Tagmentation and amplification steps were done according to manufacturer's instructions. Quality of resulting pre-enriched libraries was assessed using a high sensitivity Small DNA Fragment Analysis Kit (Agilent Technologies, DNF-477-0500). The libraries were randomly pooled per 8 or 6 samples, for the pan-cancer and three-cancer cohort, respectively, based on their relative concentrations determined using qPCR for reference genes YWHAZ, ACTB, B2M, and UBC. Each multiplex pool was concentrated to a volume of 15 µL with AMPure XP beads (Beckman Coulter Life Sciences, A63881). Finally, enrichment was performed using probes from the Illumina Exome Panel (Illumina, 20020183), probes complementary to the spike-in controls, and blocking probes against globin (anti-CEX) as previously described 22 . At the end of the enrichment workflow, equimolar library pools were prepared based on qPCR quantification with KAPA Library Quantification Kit (Roche Diagnostics, KK4854). Paired-end sequencing of both cohorts was performed (2x100 nucleotides) on a NovaSeq 6000. For the pan-cancer cohort, a NovaSeq S2 kit (Illumina, 20028315) was used with standard workflow loading of 1.55 nM (1% PhiX). For the three-cancer cohort, a NovaSeq S1 kit (Illumina, 20028318) was used with Xp workflow loading of 1.25 nM (2% PhiX). Adapter trimming and removal of reads shorter than 20 nucleotides was done with cutadapt (v1.18). Read quality was assessed with FastQC (v0.11.9) and low-quality reads were filtered: only reads where at least 80% of bases in both mates have a quality score ≥20 (99% accuracy) were kept (between 87 and 99% of reads/sample). Samples with very few reads (< 2M) were removed from all analyses. More specifically, 4 samples were removed in the pan cancer plasma cohort (1 lymphoma, 1 bladder, 1 head and neck, and 1 lung cancer sample) and 4 samples in the three-cancer cohort (3 control samples, of which 1 repeated from first cohort, and 1 ovarian cancer) as indicated in Supplemental Table 1. Quality filtered reads were then mapped with STAR 23 (v2.6.0) using the default parameters (except for --twopassMode Basic, --outFilterMatchNmin 20 and --outSAMprimaryFlag AllBestScore). The reference files for all analyses were based on genome build GRCh38 and transcriptome build Ensembl v91, complemented with spike annotations. Finally, gene counts were determined with HTSeq 24 (v0.11.0) in non-stranded mode, only considering uniquely mapping reads. Total RNA library preparation, sequencing, and quantification Total RNA sequencing libraries of the lymphoma cohort were prepared starting from 8 µL of RNA eluate using the SMARTer Stranded Total RNA-Seq Kit v3 -Pico Input Mammalian (Takara, 634487) according to the manufacturer's protocol. Equimolar library pools were prepared based on qPCR quantification with KAPA Library Quantification Kit (Roche Diagnostics, KK4854). The libraries were paired-end sequenced (2x100 nucleotides) on a NovaSeq 6000 instrument using a NovaSeq S2 kit (Illumina, 20028315) with standard workflow loading of 0.65 nM (2% PhiX). After adapter trimming with cutadapt (v1.18) and quality control with FASTQC (v0.11.9), reads were mapped with STAR 23 (v2.7.3) using default options and the GRCh38 reference described above. Resulting BAM files were deduplicated using UMI-tools (v1.0.0) based on the unique molecular identifier (UMI) sequences in the Pico v3 SMART UMI adapters. Gene counts were determined with HTSeq 24 (v0.11.0) in reverse stranded mode, only considering uniquely mapping reads. Data analyses Most data processing and visualization was done in R (v4.2.1) using tidyverse (v1.3.2) and pheatmap (v1.0.12). Other tools and R packages used for specific analyses are mentioned below. Color schemes are based on Paul Tol's technical note 25 . Detection threshold To remove noisy datapoints, we applied a threshold of at least 10 counts. This threshold was based on the median threshold that removes at least 95% of single positive genes 26 -genes that have counts in one replicate but not in the other -between control samples. MRNA concentration MRNA concentration was determined as previously described 22 . Briefly, the mass of a specific Sequin spike-in control was calculated based on the input concentration and volume of spike-in mix added to the sample. The corresponding mRNA concentration was then estimated by multiplying the spike mass by the ratio of reads mapped to the human genome versus the number of reads mapped to the specific spike, and finally dividing the obtained mass by the plasma volume of the sample. In this study, all spikes above detection threshold were considered and the geometric mean per sample was taken as the sample's mRNA concentration. Fusion gene detection FusionCatcher 27 (v1.30) was used with default settings for fusion transcript identification. Stringent filtering was applied to exclude potential false positives: transcripts with a FusionCatcher label indicative for false positives, transcripts with reads simultaneously mapping to both fusion partners, and transcripts with fusion partners less than 100 kbp apart were filtered out. Only exon-exon fusions were included and detected fusions in plasma were compared to fusions in the Fusion Gene annotation DataBase (FusionGDB) 28 , focusing on fusion genes identified in tissue samples of The Cancer Genome Atlas 29 . Differential abundance and principal component analysis DESeq2 (v1.36.0) was used for normalization and differential abundance analysis. MRNAs were pre-filtered by requiring gene counts equal to or greater than the detection threshold in at least half of the tumor or half of the control samples. In the DESeq2 result table, genes with a Benjamini-Hochberg corrected p-value (q) below 0.05 and an absolute log2 fold change above 1 were considered differentially abundant. Principal component analysis was performed with DESeq2 plotPCA on the 500 most variable genes after applying variance stabilizing transformation. Volcano plots were visualized using EnhancedVolcano (v1.14.0) and area-proportional Euler diagrams were made using eulerr (v6.1.1). Differential abundance analysis results are freely available at Zenodo (doi:10.5281/zenodo.7953707). Gene set enrichment and overrepresentation Fgsea 30 (v1.22.0) was used for preranked gene set enrichment analysis based on log2 transformed fold changes between cancer and controls obtained from DESeq2 differential abundance analysis. Significant enrichment was defined by a false discovery rate ≤ 0.05. Hallmark 9 and Canonical Pathways gene sets derived from Kyoto Encyclopedia of Genes and Genomes (KEGG) 10 , http://www.pathway.jp, were obtained via the Molecular Signatures Database, MSigDB (v2022.1) 31 . Besides these MSigDB gene sets, custom tissue and cell type specific gene sets were made using inhouse and external references as described below. Gene set enrichment analysis results and custom gene sets are available at Zenodo (doi:10.5281/zenodo.7953707). Functional analysis of mRNA sets was done using the functional annotation tool of DAVID (v2021) 32,33 and -for custom gene sets -the fora function of fgsea. For each functional analysis, a background was provided which included all mRNAs satisfying the count threshold in at least one sample. RNA Atlas cell type gene sets These cell type gene sets were defined based on total RNA sequencing data of cells and cell lines in the RNA Atlas 34 . Protein coding gene inclusion criteria were an expression of at least 16 counts after variance stabilizing transformation and an expression fold change of at least 10 compared to the cell type with second highest expression. 15 out of 32 defined cell type sets included more than 5 genes. Tabula Sapiens cell type gene sets These cell type gene sets were based on the Tabula Sapiens v1 basis matrix defined by Vorperian et al 2 . Protein coding genes with at least 5 CPM and an expression fold change of at least 10 compared to the cell type with the second highest expression were retained. 13 out of 39 defined cell type gene sets included more than 5 genes. GTEX tissue gene sets These tissue gene sets were defined using SPECS specificity scores 35 of GTEx 36 tissue samples, summarized in https://specs.cmgg.be. Criteria for protein coding gene inclusion in a certain tissue gene set were a SPECS score ≥ 0.98 and a median TPM ≥ 5. Large gene sets were reduced to the top 200 genes based on the highest SPECS score. 23 out of 27 defined tissues gene sets included more than 5 genes. HPA-RNA Atlas tissue gene sets These tissue gene sets were defined based on expression data from the Human Protein Atlas 37 (HPA, v18, https://v18.proteinatlas.org/) in combination with total RNA-sequencing data from the RNA Atlas 34 . Human Protein Atlas protein coding genes needed to have a TPM value of at least 5 and expression that is at least 10 times higher in the tissue with highest expression compared to the tissue with second highest expression. These genes sets were further finetuned by only keeping those genes with the highest expression in the same tissue in the RNA Atlas dataset. Finally, large gene sets were reduced to the top 200 genes based on the highest fold changes. 15 out of 18 defined tissue gene sets included more than 5 genes. TCGA cancer tissue gene sets These gene sets were defined similarly to the GTEX gene sets but using expression in The Cancer Genome Atlas (TCGA) cancer tissue samples, generated by the TCGA Research Network: https://www.cancer.gov/tcga. Protein coding genes with a SPECS score of at least 0.98 in a certain tissue were included in the gene set with a limit of 200 genes, based on highest SPECS score. 19 out of 23 defined cancer tissue gene sets included more than 5 genes. Cell type deconvolution NuSVR deconvolution using the Tabula Sapiens v1 basis matrix was performed as described by Vorperian et al. 2 and the accompanying GitHub repository, https://github.com/sevahn/deconvolution. As recommended by the authors, data was CPM normalized and no log transformation was performed. Note that the erythrocyte fraction is underestimated due to the absence of hemoglobin probes in mRNA capture sequencing. Tail genes To identify tail genes, transcript counts of every sample were first converted to a customized z score based on the mean and standard deviation in the reference group, including all control samples except the sample of interest. More specifically, a pseudocount was added to all normalized transcript counts followed by log2 transformation. Then, for each gene, the mean and standard deviation of these transformed counts in the reference group was used to scale the transformed counts in the sample of interest. Tail genes were then defined as genes with at least 40 counts after DESeq2 normalization and deviating more than 3 standard deviations from the mean of the control reference (|z|>3). For each gene in a sample of interest: 𝑧𝑠𝑐𝑜𝑟𝑒 !"#" = 𝑙𝑜𝑔 $ *𝑛𝑜𝑟𝑚𝑎𝑙𝑖𝑧𝑒𝑑 𝑐𝑜𝑢𝑛𝑡𝑠 !"#" + 15 -𝑟𝑒𝑓 88888 !"#" 𝑆𝐷 %"& !"#" where 𝑟𝑒𝑓 88888 !"#" and 𝑆𝐷 %"& !"#" are respectively the mean and standard deviation of log2 transformed counts for this gene over all (other) control samples. Fisher's exact tests (based on |z|>3 and donor cancer/control status) were used to select a subset of tail genes, named biomarker tail genes, that are specifically associated to the disease state. To avoid selection being biased by individual samples, Fisher's exact testing for each tail gene was done iteratively, each time leaving out a different sample, and a tail gene needed to be significant (p<0.05) in every iteration to be called a biomarker tail gene. As a result, the tail gene needs to be recurrent in at least two samples. Binary logistic regression, R glm function with binomial family, was used to classify a sample as cancer or control based on the number of biomarker tail genes. Leave-one-out cross-validation was used by iteratively leaving out a different sample from the training set and testing the model on this sample. Based on these predictions, the Receiver Operating Characteristic (ROC) curve and Area Under Curve (AUC) were calculated and visualized using pROC (v1.18.0) 38 . For internal validation, all sample labels were randomly reshuffled using the sample function in R without replacement. Next, tail gene identification and evaluation was repeated based on the new class labels (using the new control samples as reference). This process was repeated 20 times. Statistics Kruskal-Wallis tests were used to compare multiple groups, and Wilcoxon rank-sum tests were used to compare two groups. Corresponding statistical significance and effect sizes were calculated with rstatix (v0.7.2). For individual testing, p-values smaller than 0.05 were considered significant. In case of multiple testing, the Benjamini-Hochberg procedure was used to calculate false discovery rate adjusted p-values (q-values) and significance was defined as q smaller than 0.05. Significance of overlap between gene sets was determined by Fisher's exact test in GeneOverlap (v1.32.0). The universe consisted of all genes considered for the corresponding differential abundance or tail gene analyses (see filtering above). A Jaccard index was calculated to assess similarity between the gene sets, with 0% indicating no similarity and 100% indicating identical sets. + +
Data availability Raw RNA-sequencing data is available under restricted access (access can be requested) in the European Genome-phenome Archive (EGA) to preserve individuals' privacy and comply with patient consent for data sharing under the European General Data Protection Regulation. The pan-cancer and three-cancer cohort FASTQs can be found under study EGAS00001006755 (dataset EGAD00001009713). The lymphoma cohort FASTQs can be found under study EGAS00001007127 (dataset EGAD00001010259). The bladder cancer cohort FASTQs can be found under study EGAS00001003917 (dataset EGAD00001005439). Differential abundance and gene set enrichment analysis results are available (open access) in a Zenodo repository (doi:10.5281/zenodo.7953707). Code availability Code for differential abundance and tail gene analyses is available on GitHub at https://github.com/OncoRNALab/tailgenes.
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Acknowledgements Figures 1a and 5b were created with BioRender.com. We would like to thank Tim Mercer for providing the Sequin spikes, Eva Hulstaert and Francisco Avila Cobos for their support during the first phase of the project, and Anneleen Decock for her feedback on the initial draft.
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Funding This work was supported by Ghent University (BOF; BOF19/DOC/228), Ghent University Hospital, Fund for Scientific Research Flanders (FWO; 11C1623N to A.M., 11H7523N to P.D., G0B2820N), and Kom Op Tegen Kanker (Stand up to Cancer, the Flemish cancer society).
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Conflict of interest A.M., J.V. and P.M. are inventors on a patent application regarding the tail gene concept.
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diff --git a/grobid-trainer/resources/dataset/segmentation/corpus/tei/pub.1160333290.training.segmentation.tei.xml b/grobid-trainer/resources/dataset/segmentation/corpus/tei/pub.1160333290.training.segmentation.tei.xml new file mode 100644 index 0000000000..169fab8878 --- /dev/null +++ b/grobid-trainer/resources/dataset/segmentation/corpus/tei/pub.1160333290.training.segmentation.tei.xml @@ -0,0 +1,79 @@ + + + + + + + Creative Commons Non Commercial CC BY-NC: This article is distributed under the terms of the Creative Commons Attribution-NonCommercial 4.0 License (https://creativecommons.org/licenses/by-nc/4.0/) which permits non-commercial use, reproduction and distribution of the work without further permission provided the original work is attributed as specified on the SAGE and Open Access pages (https://us.sagepub.com/en-us/nam/open-access-at-sage). Journal canadien de la santé et de la maladie rénale https://doi.org/10.1177/20543581231183856 Canadian Journal of Kidney Health and Disease Volume 10: 1 -10 © The Author(s) 2023 Article reuse guidelines: sagepub.com/journals-permissions DOI: 10.1177/20543581231183856 journals.sagepub.com/home/cjk Basic Research Protocol 1183856C JKXXX10.1177/20543581231183856Canadian Journal of Kidney Health and DiseaseSharma et al research-article20232023 Basic Research Protocol: Exome Sequencing in Adults With Loin Pain Hematuria Syndrome: A Pilot Study Aditi Sharma 1 , Matthew B. Lanktree 2,3 , Sarah Liskowich 4 , Pouneh Dokouhaki 5 , and Bhanu Prasad 6,7 Abstract Background: Loin pain hematuria syndrome (LPHS) is a poorly understood clinical syndrome characterized by hematuria and either unilateral or bilateral severe kidney pain in the absence of identifiable urological disease. Loin pain hematuria syndrome imposes a significant health and economic impact with a loss of productivity and quality of life in a young population. Owing to an incomplete understanding of its pathophysiology, treatment has been limited to nonspecific pain management. Nearly 60 years after its initial description, we are no further ahead in understanding the molecular pathways involved in LPHS. Objective: To outline the study design for exome sequencing in adults with LPHS and their families. Methods: In this single-center case series, 24 patients with LPHS and 2 additional first-degree family members per participant will be recruited. DNA extracted from venous blood samples will undergo exome sequencing on the Illumina NovaSeq 6000 System at 100× depth and will be assessed for pathogenic variants in genes associated with hematuria (number of genes in: glomerular endothelium [n = 10] and basement membrane [n = 8]), and pain pathways (number of genes in: pain transduction [n = 17], conduction [n = 8], synaptic transmission [n = 37], and modulation [n = 27]). We will further examine identified potentially pathogenic variants that co-segregate with LPHS features among affected families. Conclusions: This pilot study may identify new directions for an investigation into the molecular mechanisms underlying LPHS. Abrégé Contexte: Le syndrome de lombalgie-hématurie est un syndrome clinique encore mal compris qui se caractérise par une hématurie et une forte douleur rénale unilatérale ou bilatérale en l'absence d'une maladie urologique identifiable. Le syndrome de lombalgie-hématurie a une incidence importante sur la santé et l'économie en entraînant une perte de productivité et de qualité de vie dans une population jeune. La compréhension de la physiopathologie de ce syndrome étant incomplète, le traitement a été limité à la gestion non spécifique de la douleur. Près de soixante ans après sa description initiale, nous en sommes au même point dans la compréhension des voies moléculaires impliquées dans le syndrome de lombalgie-hématurie. Objectif: Décrire le plan de l'étude pour le séquençage de l'exome chez les adultes atteints du syndrome de lombalgie-hématurie et des membres de leur famille. Méthodologie: Pour cette série de cas menée dans un seul center, nous recruterons 24 patients atteints du syndrome de lombalgie-hématurie et deux membres au premier degré de leur famille. L'ADN extrait d'échantillons de sang veineux sera soumis à un séquençage de l'exome sur le système Ilumina NovaSeq 6000 réglé à 100X de profondeur. Il sera également analysé pour la présence de variants pathogènes dans les gènes associés à l'hématurie (nombre de gènes dans l'endothélium glomérulaire [n = 10] et la membrane basale [n = 8]), et aux voies de transmission de la douleur (nombre de gènes dans la transduction [n = 17], la conduction [n = 8], la transmission synaptique [n = 37] et la modulation [n = 27] de la douleur). Nous poursuivrons l'examen des variants potentiellement pathogènes identifiés qui co-ségrègent avec les caractéristiques du syndrome de lombalgie-hématurie parmi les familles touchées. Conclusion: Cette étude pilote pourrait révéler de nouveaux axes de recherche sur les mécanismes moléculaires qui sous-tendent le syndrome de lombalgie-hématurie. Keywords loin pain hematuria syndrome, exome sequencing, rare disease, eGFR, hematuria, chronic pain Received November 9, 2022. Accepted for publication May 18, 2023. + + 2 + + Canadian Journal of Kidney Health and Disease + + What was known before Hematuria in patients with loin pain hematuria syndrome is glomerular in origin, but the source of pain is a mystery. What this adds We outline our study protocol to investigate genetic variants in the glomerular endothelium and basement membrane as potential contributors to glomerular hematuria in LPHS. We will also evaluate variants in candidate genes in pain path-ways. We anticipate that the results from this study will iden-tify new directions for an investigation into the molecular mechanisms underlying LPHS pathophysiology. Introduction Loin pain hematuria syndrome (LPHS) remains a poorly understood chronic pain disorder since its first description in 1967. 1 With a reported prevalence of .012%, 2 it typically affects young women in their late 20s or early 30s. Patients with LPHS experience debilitating flank (loin) pain, which can be unilateral or bilateral, along with gross or microscopic hematuria. 2,3 In 1996, Hebert et al proposed glomerular hematuria as the instigating event, sequentially leading to tubular obstruction, back leak of glomerular filtrate, and local parenchymal edema promoting compression of adja-cent tubules and subsequent capsular stretch leading to pain. 4 However, the absence of pain in other clinical conditions that include red blood cell and cellular cast formation including acute tubular necrosis (ATN), pyelonephritis, IgA nephropa-thy (Henoch Schoenlein Syndrome), vasculitis, and antiglo-merular basement membrane disease argues against the validity of this mechanism. We believe LPHS likely repre-sents a heterogeneous collection of cases that present with shared phenotypic features of glomerular hematuria and pain originating from the kidneys but with different pathogenesis and natural history. Glomerular hematuria in the absence of proteinuria is typically associated with defects in the glomerular filtration barrier, as most forms of podocyte injury lead to some degree of proteinuria. The remaining structural components of the filtration barrier include the glomerular basement membrane (GBM) and the specialized fenestrated glomerular endothe-lial cells (GEC). Glomerular basement membrane plays a key role in keeping the cellular elements of blood from tres-passing into the urinary space. The best technique to evaluate GBM structure is electron microscopy of a kidney biopsy. However, electron microscopy has not revealed structural deficits in the GBM structure of most patient's with LPHS. 4 There is a real need for an enhanced understanding of LPHS pathogenesis using advanced cellular and molecular technologies to explicate, detect, and diagnose both its pres-ence and probable outcome to treatment. Rapid advances in next-generation sequencing (NGS) technologies over the last 2 decades have enabled the extensive unbiased interrogation of the exome, with patient/parent trio being a successful approach for identifying candidates and de novo variants. Specifically, the trio exome analysis has added crucial insights for diseases with complex pathogenesis and a heter-ogenous rate of progressions, such as IgA nephropathy, 5 ste-roid-resistant nephrotic syndrome, 6 nephrolithiasis, and nephrocalcinosis. 7 In addition, for diseases like chronic kid-ney disease (CKD) of unknown cause, where renal ultra-sounds and kidney biopsies are uninformative and are unable to distinguish between multiple diseases, exome sequencing has led to the identification of 500 monogenic causes of CKD, 8,9 which in turn led to an increased diagnosis rate. 10 Similarly, a phenotypic spectrum of GBM abnormalities has been reported in patients with rare pathogenic variants in type IV collagen genes (COL4A3/4/5), classically associated with Alport syndrome, ranging from severe abnormalities in GBM thickness to no visible abnormalities at all. The identi-fication of more than 1000 mutations in the COL43/4/5 genes 11,12 that encode for the type IV collagen A3A4A5 het-erotrimer (a major component of the GBM), has led to the reclassification of Alport syndrome and benign familial hematuria/thin basement membrane nephropathy (TBMN) as type IV collagen nephropathies, 13 promoting genetic test-ing as the gold standard in understanding the prognosis of the phenotype. Therefore, we believe that the next logical step will be to interrogate the genes that encode the filtration bar-rier in patients with LPHS. There are 8 genes that encode GBM and receptors and 10 genes that code for endothelial cells that we know of to date. 14 While hematuria is believed to be glomerular in origin, the genesis of the pain remains an unsolved issue. The transmis-sion of pain from the viscera to the brain involves a pathway that is controlled by a series of genes that code for transduc-tion, conduction, synaptic transduction, and modulation. Specialized receptors, expressed in the peripheral termini of these neurons, allow noxious stimuli to be transduced into electrical impulses. The local membrane depolarization gen-erated by stimulus transduction is transmitted along the axon by specific channels, some of which are expressed + + 1 Dr. T. Bhanu Prasad Med Prof Corp, Regina, SK, Canada 2 Departments of Medicine and Health Research Methodology, Evidence, and Impact, McMaster University, Hamilton, ON, Canada 3 Division of Nephrology, St Joseph's Healthcare Hamilton, ON, Canada 4 Department of Academic Family Medicine, College of Medicine, University of Saskatchewan, Regina, Canada 5 Department of Pathology and Lab Medicine, University of Saskatchewan, Saskatoon, Canada 6 Section of Nephrology, Department of Medicine, Regina General Hospital, SK, Canada 7 College of Medicine, University of Saskatchewan, Regina, Canada Corresponding Author: Bhanu Prasad, Section of Nephrology, Department of Medicine, Regina General Hospital, 1440, 14th Avenue, Regina, SK S4P 0W5, Canada. Email: bprasad@sasktel.net + + Sharma et al + + 3 + + specifically in nociceptors. Transmission is modulated by specific channels, which generally act to reduce excitability. Nociceptors terminate in the spinal cord dorsal horn, forming synapses with nociceptive-specific spinal projection neurons. From the spinal cord, information is transmitted to the brain stem and then processed by a pain matrix in multiple brain regions. Descending input from the brain to the spinal cord back to the periphery can both inhibit and facilitate the trans-mission of information in nociceptive circuits. We believe that LPHS patients have either gain-of-function mutation of the pain pathways and/or a loss-of-function mutation in the pain dampening pathways. We hypothesize that rare variants identified in this study will help delineate disease pathways from a syndrome driven by clinically ascertained phenotype as a diagnosis of exclusion into one based on the molecular basis of disease that leads to hematuria and physically inca-pacitating pain (Figure 1). To improve the sensitivity and specificity to identify a responsible variant over singleton analysis, we decided to perform a family-based analysis. Exome sequencing will be performed in LPHS-affected pro-bands and participating first-degree family members, to effectively detect de novo and compound heterozygous vari-ants. In the absence of parents, analysis will be performed on the other family members such as siblings or cousins. Systematic Review of Observational Studies We searched PubMed, Embase, Scopus and Web of Science databases to identify published studies on LPHS (search term: "loin pain hematuria" OR "Loin pain-hematuria" OR "loin pain/haematuria" OR "loin pain haematuria"). The search was generated in September 2021, with no date limits set. A total of 110 articles were identified. Studies with no abstracts or articles were excluded (n = 6). Upon review of 104 LPHS articles included, encompassing 610 LPHS patients, with 68% of the articles and 87% of patients reported from the United States and United Kingdom alone. With all articles published so far focusing on symptomatic pain management using oral narcotic therapy and/or interventional management strategies such as intra-ureteric infusions, 15-17 surgical renal denervation, 18-20 radiofrequency ablation, 21-23 neuromodula-tion, 24-26 and auto-transplantation, 27-31 we did not identify any studies looking at (A) evidence of familial clustering of LPHS or its components, variability in prevalence across different ancestries, or antecedent exposures or condition, or (B) vari-ants in genes coding for proteins that are involved in GEC, GBM, or the pain pathways. Research Objectives The specific objectives of the study are as follows: Objective 1: Identify potentially pathogenic variants in 18 genes associated with hematuria 14 (Table 1). Objective 2: Identify potentially pathogenic variants in 89 genes associated with pain syndromes 32 (Table 2). Methods We plan to conduct a single-center, pilot study with the aim of decoding the molecular basis of LPHS. The study has Figure 1. Study design. + + 4 + + Canadian Journal of Kidney Health and Disease + + Table 1. Candidate Genes to be Investigated for Contributing to Hematuria in LPHS. Gene name Chromosome number Inheritance a Gene annotation Glomerular basement membrane and receptors LAMB2 chr3 AR Laminin subunit beta 2 LAMA5 chr20 AR Laminin subunit alpha 5 ITGA3 chr17 AR Integrin subunit alpha 3 COL4A3/4/5 chr2/chr2/chrX AR, AD/AR, AD/XLD Collagen type IV alpha 3/4/5 chain GPC5 chr13 Glypican 5 CD151 chr11 CD151 molecule -transmembrane 4 superfamily Endothelial cells CFH chr1 AR, AD Complement factor H CFB chr6 AR, AD, DD Complement factor B CFI chr4 AR, AD Complement factor I C3 chr19 AR, AD Complement C3 MCP chr1 AR, AD Membrane cofactor protein THBD chr20 AD Thrombomodulin PLG chr6 AR, AD Plasminogen DGKE chr17 AR Diacylglycerol kinase epsilon INF2 chr14 AD Inverted formin 2 MMACHC chr1 AR Metabolism of cobalamin associated C a Inheritance based on OMIM database: AD, autosomal dominant; AR, autosomal recessive; Mu, multifactorial; SMu, somatic mutation; XL, X linked; XLR, X linked recessive; XLD, X linked dominant Table 2. Candidate Genes to be Investigated for Understanding Pain in LPHS Patients. Gene name Chromosome number Inheritance a Gene annotation Pain transduction TRPV1/2/3/4 chr17/chr17/chr17/chr12 TRPV3/4: AD Transient receptor potential cation channel subfamily V member 1/2/3/4 P2RX3 chr11 P2X purinergic receptor TRPM8 chr2 Transient receptor potential cation channel subfamily M member 8 TRPA1 chr8 AD Transient receptor potential cation channel subfamily A member 1 P2RY12 chr3 AR Purinergic receptor P2y12 BDKRB1/2 chr14/chr14 Bradykinin receptor B1/2 HTR3a chr11 5-Hydroxytryptamine receptor 3a ACCN1/2 chr17/chr12 Acid sensing ion channel subunit 1/2 TRPC/P chr13 Transient receptor potential canonical Pain conduction SCN10A chr3 AD Sodium voltage-gated channel alpha subunit 10 SCN11A chr3 AD Sodium voltage-gated channel alpha subunit 11 SCN1,3,8A chr2/chr2/chr12 AD/AD/AD Sodium channel protein type 1/3/8A SCN9A chr2 AR, AD Sodium voltage-gated channel alpha subunit 9 KCNQ chr11 AR, AD Potassium voltage-gated channel subfamily Q Pain synaptic transmission NR1, 2 chr9/ AR, AD Nuclear receptor subfamily 1/2 GRIA1-4 chr5/chr4/chrX/chr11 AR, AD/ AD/ XLR/ AD AMPA receptor 1-4 NK1R chr2 Tachykinin receptor 1 CACNA1A-I, S chr19/chr9/chr12/chr3/ chr1/chrX/chr17/chr16/ chr22/chr1 AD/ AR/ AD/ AR, AD/ AD/ XL, XLR/ AD/ AD/ AD/ A R, AD Calcium voltage-gated channel subunit alpha1 A-S CACNA2D1 chr7 AR Calcium voltage-gated channel auxiliary subunit alpha2 delta 1 (continued) + + Sharma et al + + 5 + + Gene name Chromosome number Inheritance a Gene annotation Pain modulation BDNF chr11 Brain-derived neurotrophic factor OPRD1/M1/K1 chr1/ chr6/ chr8 Opioid receptor delta 1/Mu1/kappa 1 CNR1 chr6 Cannabinoid receptor 1 GABRs (GABRD/E/ P/Q/A1-A6/BR1-BR3/G1-G3/R1-R3 chr1/chrX/chr5/chrX/chr5/ chr4/chrX/chr4/chr15/ chr5/chr4/chr5/chr15/ chr4/chr5/chr15/chr6/ chr6/chr3 AD/ M/ M/ M/ AD/ AD, Mu/ XL/ M/ AD/ M/ AD/AD/ AD/ M/ AD/ M/ M/ M/ M Gamma-aminobutyric acid type A receptors TNF chr6 AD Tumor necrosis factor PLA2G2A chr1 AD, SMu Phospholipase A2 IL1/6/12/18 chr2/chr7/chr3/chr11 M/ Mu, SMu, AR, AD/ M/ M Interleukin 1/6/12/18 COX2 chrMT Cytochrome C oxidase subunit 2 NTRK1 chr1 AR High affinity nerve growth factor receptor NGF chr1 AR Beta-nerve growth factor GDNF chr5 AD Glial cell line-derived neurotrophic factor LIF chr22 AD Leukemia inhibitory factor CCL2 chr17 C-C motif chemokine 2 CNR2 chr1 Cannabinoid receptor 2 TLR2/4 chr4/chr9 AD, SMu/ M Toll-like receptor 2/4 P2RX4/7 chr12/chr12 P2X purinoceptor 4/7 CX3CR1 chr3 CX3C chemokine receptor 1 a Inheritance based on OMIM database: AD, autosomal dominant; AR, autosomal recessive; Mu, multifactorial; SMu, somatic mutation; XL, X linked; XLR, X linked recessive; M, missing or no information in OMIM database. Table 2. (continued) been approved by the Saskatchewan Health Authority Research Ethics Board (REB-22-66). Design Number of Subjects Twenty-four LPHS patients referred to the nephrology clinic (run by the corresponding author) were approached by the study coordinator, and all 24 agreed to participate in the study. Recruitment of 2 family members of the LPHS patient (parents/siblings/cousins/children) is ongoing. We are the first study to identify and report 2 patients with LPHS (12, 17) with a positive family history of LPHS (Figure 2). In addition, 6 LPHS patients (LPHS: 01, 02, 04, 05, 11, and 21) had a positive family history of intermittent hematuria (Figure 2). Duration of Study Period The study period extended for about 3 years (2022-2025). Participant Selection and Informed Consent The LPHS patients have been identified by BP (Nephrologist) and FG (Urologist), based on the inclusion and exclusion cri-teria defined by Spetie et al 4 All patients and family mem-bers that meet the eligibility criteria will need to complete the written consent forms followed by baseline data collec-tion (detailed below). Inclusion criteria for LPHS patient • ≥18 years of age • Patients diagnosed with LPHS (by nephrologist and/ or urologist). Exclusion criteria for LPHS patient • Urological causes of flank pain and hematuria (obstructive uropathy, nephrolithiasis, pyelonephritis, polycystic kidney disease, renal artery embolism, renal artery dissection, renal papillary necrosis, renal vein thrombosis, left renal vein entrapment [nut-cracker syndrome], renal trauma, or renal tumor). • Additional functional/structural reason for hematuria by kidney biopsy or other interventions. Inclusion/exclusion criteria for family members • Only family members (parents/siblings/children/first cousin) of the patient who provide informed consent. Data Collection, Sample Collection, and Exome Analysis Demographic Data Data related to age, sex, gender, self-reported race/ethnic-ity, occupation, socioeconomic status, weight, height, + + 6 + + Canadian Journal of Kidney Health and Disease + + Figure 2. Pedigree of LPHS patients with a family history of LPHS or isolated hematuria (IH). Roman numbers represent the generations of the family. Note. Proband is indicated by arrow. Abbreviations: k, kidney stones; m, migraine. comorbidities, family history of diseases (eg, kidney stones, isolated hematuria, LPHS, diabetes, hypertension, chronic fatigue syndrome, mood disorders, asthma, and allergies), the location of pain (bilateral or unilateral), the number and frequency of pain medications, and pain score using the brief pain inventory form and PainDetect questionnaire will be collected for the LPHS patients. Laboratory Data Complete blood count, serum electrolytes, serum urea and creatinine, albumin creatinine ratio, and urine analysis. Genetic Data Rare variants in genes associated with hematuria (n = 18) and in pain transduction (n = 17), conduction (n = 8), syn-aptic transmission (n = 37), and pain modulation (n = 27). Hierarchical Clustering of Patient Phenotype Data Patients will be grouped based on their demographic, labora-tory, and genetic data using unsupervised hierarchical clus-tering (Ward's method with Euclidean distances). 33 The data will be presented as clustered dendrograms. Sample Collection, Storage, and Preparation Venous blood samples will be collected for study probands and participating family members in PAXgene Blood RNA Tubes (Qiagen, Hilden, Germany). DNA will be extracted from PAXgene Blood RNA Tube using the New England Biolabs Monarch Genomic DNA Purification Kit as per the protocol by Kruhøffer et al. 34 DNA libraries will be con-structed using the Illumina DNA Prep with Enrichment, Tagmentation kit and IDT xGen Exome Research Panel v2 with xGen Universal Blockers-NXT Mix and dual unique + + Sharma et al + + 7 + + barcodes. Paired-end sequencing (2 × 150 bps) on the Illumina NovaSeq 6000 System at 100× depth for exome sequencing. Library preparation and sequencing will be per-formed for all family members at the same time to minimize potential artifactual differences due to sample preparation. DNA samples will be stored at -80° centigrade to allow for future verification studies. Exome Analysis As depicted in Figure 3, raw fastq files will be processed to remove adapter sequences using cutadapt (v1.11). 35 Reads with quality score of ≥30 will be retained for analysis. Contaminating reads will be removed after aligning of the processed reads to the human genome using BWA 36 and Samtools. 37 The reads will be preprocessed using the Genomic Analysis Tool Kit (GATK), as per the recommen-dations in the Best Practices Workflow by the GATK 38 for all positions with ≥20× coverage, genotype quality ≥20, and minor read ratio ≥0.2 for indel alignment, base quality score recalibration, base alignment quality scoring, and variant calling (single-nucleotide variants [SNVs], indels, short tan-dem repeats [STRs], structural variant [SV]). Variant allele frequency (VAF) will be calculated as the percentage of sequence reads observed for the alternative allele compared to all coverage of that nucleotide. Exome data will first be evaluated for genes associated with isolated hematuria and pain (as listed in Tables 1 and 2). The variants will be further filtered and prioritized using VarSeq software (Golden Helix, Bozeman, MT, USA) and variant effect predictor. Benign variants with MAF (minor allele frequency) >1% in any ancestry will be eliminated using the 1000 Genomes Project, 39 dbSNP, 40 gnomAD 41 and National Institutes of Health (NIH)'s All of Us. 42 In silico bioinformatic prediction of pathogenicity of variants will be performed using the fol-lowing prediction algorithms: scale-invariant feature trans-form (SIFT), 43 PolyPhen2, 44 Mutation Taster, 45 CADD, 46 VEST, 47 and FATHMM. 48 Finally, the candidate variants will be checked against the human gene mutation database (HGMD) 41 , LOVD, 12 Clinvar, 49 and Alport database. 50 A multidisciplinary team will then review each variant for evi-dence of pathogenicity and contribution to the phenotype and classify them according to the American College of Medical Genetics guidelines. 51 Possible pathogenic loci will be screened according to 3 heredity models, namely autoso-mal recessive (AR) inheritance, autosomal dominant (AD), and X-linked inheritance. Sanger sequencing will be carried out to validate potentially pathogenic variants identified through high-throughput exome sequencing. Special atten-tion will be paid to de novo variants, not present in parents, as well as variants with variant allele frequencies suggestive of potential somatic variation. Finally, we will also assess for the presence of copy number variation using CoNIFER, 52 cn.MOPS, 53 and CNVkit. 54 To evaluate the pathogenicity of a rare variant, we will look at the segregation of variants among all sequenced family members using (A) probability-based models by Helbig et al 55 and Jarvik et al, 56 and (B) gene-based segregation methods. 57 Potential Risks to the Participants In Canada, people are protected from being required to provide the results of a genetic test by the Genetic Non-Discrimination Act. 58 The genetic results will not be Figure 3. Pipeline for exome data analysis. + + 8 + + Canadian Journal of Kidney Health and Disease + + disclosed to any third party such as employers, insurance companies, or educational institutions. The confidentiality of the participant will be respected. All collected samples will be assigned a unique study number, with no reference to indi-vidual identifiers. As this research involves looking at genetic information, it carries the risk of identifying an underlying genetic change(s) which are unrelated to this study and have the potential of affecting the participant. However, this research is being conducted for the scientific purpose of understanding only the cause of pain and hematuria in LPHS patients. In addition, the results of this research project will not be placed in the participant's medical record. All efforts will be made to safeguard participants' privacy. + +
Author Contributions BP conceived and designed the study. AS wrote the initial draft. ML and SL assisted with the drafts. ML provided advice regarding genetic study design. BP edited the final manuscript. All authors read and approved the final manuscript. Declaration of Conflicting Interests The author(s) declared the following potential conflicts of interest with respect to the research, authorship, and/or publication of this article: BP has received speaker and advisory fees from Bayer, Otsuka and Astra Zeneca. MBL has received speaker and advisory fees from Otsuka, Reata, Bayer, and Sanofi Genzyme.
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Funding The author(s) disclosed receipt of the following financial support for the research, authorship, and/or publication of this article: The author(s) received financial support for the research, authorship, and/or publication of this article from the Hospitals of Regina Foundation. MBL has funding support from CIHR project grant (grant no. 201909-PJT). The funding sources had no role in the design, conduct, and analysis of the study or in the decision to sub-mit the manuscript for publication.
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Ethics Approval and Consent to Participate The study will be conducted in accordance with the second edition of the Tri-Council Policy Statement-Ethical Conduct for Research Involving Humans-TCPS 2. We have received REB certificate of approval for the study (REB-22-66). Written informed consent will be obtained from all the participants in the study. Consent for Publication Not applicable as there is no patient identifying information in this article.
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Availability of Data and Materials The data sets used and/or analyzed during the current study are available from the corresponding author on reasonable request.
+ + ORCID iDs Matthew B. Lanktree https://orcid.org/0000-0002-5750-6286 Bhanu Prasad https://orcid.org/0000-0002-1139-4821 + + References 1. Little PJ, Sloper JS, de Wardener HE. A syndrome of loin pain and haematuria associated with disease of peripheral renal arteries. Q J Med. 1967;36(142):253-259 2. Vakili STT, Alam T, Sollinger H. Loin pain hematuria syn-drome. Am J Kidney Dis. 2014;64(3):460-472. doi:10.1053/j. ajkd.2014.01.439 3. Aber GM, Higgins PM. The natural history and management of the loin pain/haematuria syndrome. Br J Urol. 1982;54(6):613-615. doi:10.1111/j.1464-410x.1982.tb13607.x 4. Spetie DN, Nadasdy T, Nadasdy G, et al. Proposed pathogen-esis of idiopathic loin pain-hematuria syndrome. Am J Kidney Dis. 2006;47(3):419-427. doi:10.1053/j.ajkd.2005.11.029 5. Cox SN, Pesce F, El-Sayed Moustafa JS, et al. Multiple rare genetic variants co-segregating with familial IgA nephropathy all act within a single immune-related network. 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