0 10 Functional functional JJ 11 19 analysis analysis NN 20 22 of of IN 23 33 cis-linked cis-linked JJ 34 44 regulatory regulatory JJ 45 54 sequences sequence NNS 55 57 in in IN 58 61 the the DT 62 65 HLA HLA NNP 66 69 DRA DRA NNP 70 78 promoter promoter NN 79 81 by by IN 82 95 transcription transcription NN 96 98 in in FW 99 104 vitro vitro FW 104 105 . . . 107 110 Two two CD 111 120 consensus consensus NN 121 130 sequences sequence NNS 130 131 , , , 132 138 called call VBN 139 140 X X NNP 141 144 and and CC 145 146 Y Y NNP 147 152 boxes box NNS 152 153 , , , 154 161 capable capable JJ 162 164 of of IN 165 172 binding bind VBG 173 180 nuclear nuclear JJ 181 189 proteins protein NNS 190 193 and and CC 194 204 regulating regulate VBG 205 215 expression expression NN 216 218 in in IN 219 220 B B NNP 221 226 cells cell NNS 227 231 have have VBP 232 236 been be VBN 237 244 defined define VBN 245 251 within within IN 252 255 the the DT 256 265 immediate immediate JJ 266 274 upstream upstream JJ 275 281 region region NN 282 284 of of IN 285 290 major major JJ 291 309 histocompatibility histocompatibility NN 310 317 complex complex NN 318 319 ( ( ( 319 322 MHC MHC NNP 322 323 ) ) ) 324 329 class class NN 330 332 II ii CD 333 342 promoters promoter NNS 342 343 . . . 344 350 Unlike unlike IN 351 356 other other JJ 357 362 class class NN 363 365 II ii CD 366 375 promoters promoter NNS 375 376 , , , 377 380 the the DT 381 387 HLA-DR HLA-DR NNP 388 393 alpha alpha NN 394 395 ( ( ( 395 398 DRA DRA NNP 398 399 ) ) ) 400 408 promoter promoter NN 409 413 also also RB 414 422 contains contain VBZ 423 426 one one CD 427 434 element element NN 435 444 identical identical JJ 445 447 to to TO 448 451 the the DT 452 453 “ `` `` 453 460 octamer octamer NN 460 461 ” '' '' 462 467 motif motif NN 468 470 of of IN 471 485 immunoglobulin immunoglobulin NN 486 494 variable variable JJ 495 501 region region NN 502 511 promoters promoter NNS 512 516 that that WDT 517 519 is be VBZ 520 531 responsible responsible JJ 532 535 for for IN 536 537 B B NNP 538 551 cell-specific cell-specific JJ 552 565 transcription transcription NN 565 566 . . . 567 571 This this DT 572 573 “ `` `` 573 580 octamer octamer NN 580 581 ” '' '' 582 584 in in IN 585 588 the the DT 589 596 context context NN 597 599 of of IN 600 603 DRA DRA NNP 604 611 appears appear VBZ 612 619 capable capable JJ 620 622 of of IN 623 630 binding bind VBG 631 635 both both CC 636 639 the the DT 640 650 ubiquitous ubiquitous JJ 651 652 ( ( ( 652 657 OTF-1 otf-1 NN 657 658 ) ) ) 659 662 and and CC 663 680 lymphoid-specific lymphoid-specific JJ 681 682 ( ( ( 682 687 OTF-2 otf-2 NN 687 688 ) ) ) 689 690 “ `` `` 690 697 octamer octamer NN 697 698 ” '' '' 699 706 binding binding NN 707 715 proteins protein NNS 715 716 , , , 717 720 but but CC 721 723 at at IN 724 729 least least JJS 730 733 one one CD 734 739 other other JJ 740 748 distinct distinct JJ 749 750 “ `` `` 750 757 octamer octamer NN 757 758 ” '' '' 759 766 complex complex NN 767 770 was be VBD 771 776 found find VBN 776 777 . . . 778 780 In in IN 781 786 order order NN 787 789 to to TO 790 802 characterize characterize VB 803 806 the the DT 807 815 function function NN 816 818 of of IN 819 829 cis-acting cis-acting JJ 830 838 elements element NNS 838 839 , , , 840 842 we we PRP 843 847 have have VBP 848 857 developed develop VBN 858 860 an an DT 861 863 in in FW 864 869 vitro vitro FW 870 876 system system NN 877 879 in in IN 880 885 which which WDT 886 887 a a DT 888 891 DRA DRA NNP 892 900 promoter promoter NN 901 910 construct construct NN 911 913 is be VBZ 914 925 transcribed transcribe VBN 926 930 more more RBR 931 942 efficiently efficiently RB 943 945 in in IN 946 954 extracts extract NNS 955 959 from from IN 960 961 B B NNP 962 967 cells cell NNS 968 972 than than IN 973 975 in in IN 976 984 extracts extract NNS 985 989 from from IN 990 995 class class NN 996 1007 II-negative ii-negative JJ 1008 1012 HeLa HeLa NNP 1013 1018 cells cell NNS 1018 1019 . . . 1020 1021 5 5 CD 1021 1022 ’ ' SYM 1023 1031 deletion deletion NN 1032 1042 constructs construct NNS 1043 1048 which which WDT 1049 1055 lacked lack VBD 1056 1059 the the DT 1060 1061 Y y NN 1062 1065 box box NN 1065 1066 , , , 1067 1070 but but CC 1071 1079 retained retain VBD 1080 1083 the the DT 1084 1085 “ `` `` 1085 1092 octamer octamer NN 1092 1093 ” '' '' 1094 1099 motif motif NN 1100 1103 and and CC 1104 1108 TATA TATA NNP 1109 1112 box box NN 1113 1117 were be VBD 1118 1128 completely completely RB 1129 1137 inactive inactive JJ 1137 1138 , , , 1139 1142 and and CC 1143 1151 internal internal JJ 1152 1160 deletion deletion NN 1161 1163 of of IN 1164 1167 the the DT 1168 1169 Y Y NNP 1170 1173 box box NN 1174 1181 reduced reduce VBD 1182 1195 transcription transcription NN 1196 1198 by by IN 1199 1201 95 95 CD 1201 1202 % % NN 1202 1203 . . . 1204 1209 Using use VBG 1210 1221 supercoiled supercoile VBN 1221 1222 , , , 1223 1226 but but CC 1227 1230 not not RB 1231 1237 linear linear JJ 1238 1247 templates template NNS 1247 1248 , , , 1249 1251 we we PRP 1252 1260 observed observe VBD 1261 1272 differences difference NNS 1273 1275 in in IN 1276 1289 transcription transcription NN 1290 1302 efficiencies efficiency NNS 1303 1307 from from IN 1308 1317 templates template NNS 1318 1325 lacking lack VBG 1326 1328 or or CC 1329 1339 disrupting disrupt VBG 1340 1343 the the DT 1344 1345 X X NNP 1346 1355 consensus consensus NN 1356 1363 element element NN 1364 1368 that that WDT 1369 1376 reflect reflect VBP 1377 1384 effects effect NNS 1385 1387 of of IN 1388 1394 random random JJ 1395 1406 replacement replacement NN 1407 1409 of of IN 1410 1411 X X NNP 1412 1415 box box NN 1416 1425 sequences sequence NNS 1426 1428 in in IN 1429 1438 transient transient JJ 1439 1449 expression expression NN 1450 1456 assays assay NNS 1456 1457 . . . 1458 1471 Demonstration demonstration NN 1472 1474 of of IN 1475 1478 the the DT 1479 1487 complete complete JJ 1488 1498 dependence dependence NN 1499 1501 on on IN 1502 1505 the the DT 1506 1507 Y y NN 1508 1511 box box NN 1512 1514 in in IN 1515 1519 this this DT 1520 1526 system system NN 1527 1535 suggests suggest VBZ 1536 1540 that that IN 1540 1541 , , , 1542 1549 despite despite IN 1550 1553 its its PRP$ 1554 1566 demonstrated demonstrate VBN 1567 1577 importance importance NN 1578 1580 in in IN 1581 1584 the the DT 1585 1588 DRA DRA NNP 1589 1597 promoter promoter NN 1597 1598 , , , 1599 1602 the the DT 1603 1606 DRA DRA NNP 1607 1608 “ `` `` 1608 1615 octamer octamer NN 1615 1616 ” '' '' 1617 1621 does do VBZ 1622 1625 not not RB 1626 1633 utilize utilize VB 1634 1639 OTF-2 otf-2 NN 1640 1642 in in IN 1643 1644 a a DT 1645 1651 manner manner NN 1652 1661 analogous analogous JJ 1662 1664 to to TO 1665 1679 immunoglobulin immunoglobulin NN 1680 1689 promoters promoter NNS 1690 1692 in in IN 1693 1694 B B NNP 1695 1700 cells cell NNS 1700 1701 . . .