0 11 Synergistic synergistic JJ 12 24 interactions interaction NNS 25 32 between between IN 33 44 overlapping overlap VBG 45 52 binding bind VBG 53 58 sites site NNS 59 62 for for IN 63 66 the the DT 67 72 serum serum NN 73 81 response response NN 82 88 factor factor NN 89 92 and and CC 93 98 ELK-1 elk-1 NN 99 107 proteins protein NNS 108 115 mediate mediate VBP 116 120 both both CC 121 126 basal basal JJ 127 138 enhancement enhancement NN 139 142 and and CC 143 150 phorbol phorbol NN 151 156 ester ester NN 157 171 responsiveness responsiveness NN 172 174 of of IN 175 182 primate primate JJ 183 198 cytomegalovirus cytomegalovirus NN 199 204 major major JJ 205 220 immediate-early immediate-early JJ 221 230 promoters promoter NNS 231 233 in in IN 234 242 monocyte monocyte NN 243 246 and and CC 247 259 T-lymphocyte T-lymphocyte NNP 260 264 cell cell NN 265 270 types type NNS 270 271 . . . 273 288 Cytomegalovirus Cytomegalovirus NNP 289 290 ( ( ( 290 293 CMV CMV NNP 293 294 ) ) ) 295 304 infection infection NN 305 307 is be VBZ 308 321 nonpermissive nonpermissive JJ 322 324 or or CC 325 335 persistent persistent JJ 336 338 in in IN 339 343 many many JJ 344 352 lymphoid lymphoid JJ 353 356 and and CC 357 364 myeloid myeloid JJ 365 369 cell cell NN 370 375 types type NNS 376 379 but but CC 380 383 can can MD 384 386 be be VB 387 396 activated activate VBN 397 399 in in IN 400 414 differentiated differentiate VBN 415 426 macrophages macrophage NNS 426 427 . . . 428 430 We we PRP 431 435 have have VBP 436 441 shown show VBN 442 451 elsewhere elsewhere RB 452 456 that that IN 457 461 both both CC 462 465 the the DT 466 471 major major JJ 472 487 immediate-early immediate-early JJ 488 492 gene gene NN 493 494 ( ( ( 494 497 MIE MIE NNP 497 498 ) ) ) 499 502 and and CC 503 508 lytic lytic JJ 509 514 cycle cycle NN 515 525 infectious infectious JJ 526 533 progeny progeny NN 534 539 virus virus NN 540 550 expression expression NN 551 554 can can MD 555 557 be be VB 558 565 induced induce VBN 566 568 in in IN 569 578 otherwise otherwise RB 579 592 nonpermissive nonpermissive JJ 593 606 monocyte-like monocyte-like JJ 607 612 U-937 u-937 NN 613 617 cell cell NN 618 626 cultures culture NNS 627 635 infected infect VBN 636 640 with with IN 641 647 either either CC 648 653 human human JJ 654 657 CMV CMV NNP 658 659 ( ( ( 659 663 HCMV HCMV NNP 663 664 ) ) ) 665 667 or or CC 668 674 simian simian JJ 675 678 CMV CMV NNP 679 680 ( ( ( 680 684 SCMV SCMV NNP 684 685 ) ) ) 686 688 by by IN 689 698 treatment treatment NN 699 703 with with IN 704 707 the the DT 708 715 phorbol phorbol NN 716 721 ester ester NN 722 758 12-O-tetradecanoylphorbol-13-acetate 12-O-tetradecanoylphorbol-13-acetate NNP 759 760 ( ( ( 760 763 TPA TPA NNP 763 764 ) ) ) 764 765 . . . 766 769 Two two CD 770 779 multicopy multicopy JJ 780 785 basal basal JJ 786 794 enhancer enhancer NN 795 801 motifs motif NNS 802 808 within within IN 809 812 the the DT 813 817 SCMV SCMV NNP 818 821 MIE MIE NNP 822 830 enhancer enhancer NN 830 831 , , , 832 838 namely namely RB 838 839 , , , 840 842 11 11 CD 843 849 copies copy NNS 850 852 of of IN 853 856 the the DT 857 862 16-bp 16-bp JJ 863 869 cyclic cyclic JJ 870 873 AMP AMP NNP 874 882 response response NN 883 890 element element NN 891 892 ( ( ( 892 895 CRE CRE NNP 895 896 ) ) ) 897 900 and and CC 901 902 3 3 CD 903 909 copies copy NNS 910 912 of of IN 913 918 novel novel JJ 919 924 17-bp 17-bp JJ 925 930 serum serum NN 931 939 response response NN 940 946 factor factor NN 947 948 ( ( ( 948 951 SRF SRF NNP 951 952 ) ) ) 953 960 binding binding NN 961 966 sites site NNS 967 975 referred refer VBN 976 978 to to TO 979 981 as as IN 982 985 the the DT 986 989 SNE SNE NNP 990 991 ( ( ( 991 1008 SRF/NFkappaB-like srf/nfkappab-like JJ 1009 1016 element element NN 1016 1017 ) ) ) 1017 1018 , , , 1019 1021 as as RB 1022 1026 well well RB 1027 1029 as as IN 1030 1034 four four CD 1035 1044 classical classical JJ 1045 1053 NFkappaB NFkappaB NNP 1054 1059 sites site NNS 1060 1066 within within IN 1067 1070 the the DT 1071 1075 HCMV HCMV NNP 1076 1083 version version NN 1083 1084 , , , 1085 1095 contribute contribute VBP 1096 1098 to to TO 1099 1102 TPA TPA NNP 1103 1117 responsiveness responsiveness NN 1118 1120 in in IN 1121 1130 transient transient JJ 1131 1137 assays assay NNS 1138 1140 in in IN 1141 1149 monocyte monocyte NN 1150 1153 and and CC 1154 1160 T-cell t-cell NN 1161 1166 types type NNS 1166 1167 . . . 1168 1171 The the DT 1172 1176 SCMV SCMV NNP 1177 1180 SNE SNE NNP 1181 1186 sites site NNS 1187 1194 contain contain VBP 1195 1204 potential potential JJ 1205 1216 overlapping overlap VBG 1217 1221 core core NN 1222 1233 recognition recognition NN 1234 1241 binding binding NN 1242 1248 motifs motif NNS 1249 1252 for for IN 1253 1256 SRF SRF NNP 1256 1257 , , , 1258 1270 Rel/NFkappaB Rel/NFkappaB NNP 1270 1271 , , , 1272 1275 ETS ETS NNP 1275 1276 , , , 1277 1280 and and CC 1281 1284 YY1 yy1 NN 1285 1290 class class NN 1291 1304 transcription transcription NN 1305 1312 factors factor NNS 1313 1316 but but CC 1317 1321 fail fail VBP 1322 1324 to to TO 1325 1332 respond respond VB 1333 1335 to to TO 1336 1342 either either CC 1343 1348 serum serum NN 1349 1351 or or CC 1352 1357 tumor tumor NN 1358 1366 necrosis necrosis NN 1367 1373 factor factor NN 1374 1379 alpha alpha NN 1379 1380 . . . 1381 1390 Therefore therefore RB 1390 1391 , , , 1392 1394 to to TO 1395 1403 evaluate evaluate VB 1404 1407 the the DT 1408 1417 mechanism mechanism NN 1418 1420 of of IN 1421 1424 TPA TPA NNP 1425 1439 responsiveness responsiveness NN 1440 1442 of of IN 1443 1446 the the DT 1447 1450 SNE sne NN 1451 1457 motifs motif NNS 1458 1461 and and CC 1462 1464 of of IN 1465 1466 a a DT 1467 1474 related related JJ 1475 1480 16-bp 16-bp JJ 1481 1484 SEE see NN 1485 1486 ( ( ( 1486 1493 SRF/ETS srf/ets NN 1494 1501 element element NN 1501 1502 ) ) ) 1503 1508 motif motif NN 1509 1514 found find VBN 1515 1517 in in IN 1518 1521 the the DT 1522 1526 HCMV HCMV NNP 1527 1530 and and CC 1531 1541 chimpanzee chimpanzee NN 1542 1545 CMV CMV NNP 1546 1549 MIE MIE NNP 1550 1559 enhancers enhancer NNS 1559 1560 , , , 1561 1563 we we PRP 1564 1568 have have VBP 1569 1577 examined examine VBN 1578 1581 the the DT 1582 1592 functional functional JJ 1593 1602 responses response NNS 1603 1606 and and CC 1607 1614 protein protein NN 1615 1622 binding binding NN 1623 1633 properties property NNS 1634 1636 of of IN 1637 1649 multimerized multimerized JJ 1650 1659 wild-type wild-type JJ 1660 1663 and and CC 1664 1670 mutant mutant JJ 1671 1679 elements element NNS 1680 1685 added add VBN 1686 1694 upstream upstream RB 1695 1697 to to TO 1698 1701 the the DT 1702 1706 SCMV SCMV NNP 1707 1710 MIE MIE NNP 1711 1713 or or CC 1714 1720 simian simian JJ 1721 1726 virus virus NN 1727 1729 40 40 CD 1730 1737 minimal minimal JJ 1738 1746 promoter promoter NN 1747 1754 regions region NNS 1755 1757 in in IN 1758 1761 the the DT 1762 1767 U-937 u-937 NN 1767 1768 , , , 1769 1774 K-562 K-562 NNP 1774 1775 , , , 1776 1781 HL-60 HL-60 NNP 1781 1782 , , , 1783 1788 THP-1 thp-1 NN 1788 1789 , , , 1790 1793 and and CC 1794 1800 Jurkat Jurkat NNP 1801 1805 cell cell NN 1806 1811 lines line NNS 1811 1812 . . . 1813 1819 Unlike unlike IN 1820 1829 classical classical JJ 1830 1838 NFkappaB NFkappaB NNP 1839 1844 sites site NNS 1844 1845 , , , 1846 1853 neither neither CC 1854 1857 the the DT 1858 1861 SNE SNE NNP 1862 1865 nor nor CC 1866 1869 the the DT 1870 1873 SEE see NN 1874 1879 motif motif NN 1880 1889 responded respond VBD 1890 1892 to to TO 1893 1904 phosphatase phosphatase NN 1905 1915 inhibition inhibition NN 1916 1918 by by IN 1919 1926 okadaic okadaic JJ 1927 1931 acid acid NN 1931 1932 . . . 1933 1940 However however RB 1940 1941 , , , 1942 1945 the the DT 1946 1949 TPA TPA NNP 1950 1964 responsiveness responsiveness NN 1965 1967 of of IN 1968 1972 both both DT 1973 1976 CMV CMV NNP 1977 1985 elements element NNS 1986 1992 proved prove VBD 1993 1995 to to TO 1996 2003 involve involve VB 2004 2015 synergistic synergistic JJ 2016 2028 interactions interaction NNS 2029 2036 between between IN 2037 2040 the the DT 2041 2045 core core NN 2046 2049 SRF srf NN 2050 2057 binding binding NN 2058 2062 site site NN 2063 2064 ( ( ( 2064 2074 CCATATATGG CCATATATGG NNP 2074 2075 ) ) ) 2076 2079 and and CC 2080 2083 the the DT 2084 2092 adjacent adjacent JJ 2093 2101 inverted invert VBN 2102 2105 ETS ets NN 2106 2113 binding binding NN 2114 2120 motifs motif NNS 2121 2122 ( ( ( 2122 2126 TTCC TTCC NNP 2126 2127 ) ) ) 2127 2128 , , , 2129 2134 which which WDT 2135 2145 correlated correlate VBD 2146 2154 directly directly RB 2155 2159 with with IN 2160 2169 formation formation NN 2170 2172 of of IN 2173 2174 a a DT 2175 2180 bound bind VBN 2181 2191 tripartite tripartite JJ 2192 2199 complex complex NN 2200 2210 containing contain VBG 2211 2215 both both CC 2216 2219 the the DT 2220 2228 cellular cellular JJ 2229 2232 SRF srf NN 2233 2236 and and CC 2237 2242 ELK-1 elk-1 NN 2243 2251 proteins protein NNS 2251 2252 . . . 2253 2257 This this DT 2258 2265 protein protein NN 2266 2273 complex complex NN 2274 2277 was be VBD 2278 2282 more more RBR 2283 2291 abundant abundant JJ 2292 2294 in in IN 2295 2300 U-937 U-937 NNP 2300 2301 , , , 2302 2307 K-562 K-562 NNP 2307 2308 , , , 2309 2312 and and CC 2313 2317 HeLa HeLa NNP 2318 2322 cell cell NN 2323 2331 extracts extract NNS 2332 2336 than than IN 2337 2339 in in IN 2340 2344 Raji Raji NNP 2344 2345 , , , 2346 2348 HF HF NNP 2348 2349 , , , 2350 2356 BALB/c BALB/c NNP 2357 2360 3T3 3t3 NN 2360 2361 , , , 2362 2364 or or CC 2365 2370 HL-60 hl-60 NN 2371 2376 cells cell NNS 2376 2377 , , , 2378 2381 but but CC 2382 2385 the the DT 2386 2393 binding binding NN 2394 2402 activity activity NN 2403 2406 was be VBD 2407 2414 altered alter VBN 2415 2419 only only RB 2420 2427 twofold twofold RB 2428 2433 after after IN 2434 2437 TPA TPA NNP 2438 2447 treatment treatment NN 2447 2448 . . . 2449 2450 A a DT 2451 2458 40-fold 40-fold JJ 2459 2470 stimulation stimulation NN 2471 2473 of of IN 2474 2489 chloramphenicol chloramphenicol NN 2490 2507 acetyltransferase acetyltransferase NN 2508 2516 activity activity NN 2517 2525 mediated mediate VBN 2526 2528 by by IN 2529 2533 four four CD 2534 2540 tandem tandem JJ 2541 2548 repeats repeat NNS 2549 2551 of of IN 2552 2555 the the DT 2556 2559 SNE sne NN 2560 2565 could could MD 2566 2568 be be VB 2569 2576 induced induce VBN 2577 2583 within within IN 2584 2585 2 2 CD 2586 2587 h h NN 2588 2589 ( ( ( 2589 2592 and and CC 2593 2595 up up RB 2596 2598 to to TO 2599 2607 250-fold 250-fold JJ 2608 2614 within within IN 2615 2616 6 6 CD 2617 2618 h h NN 2618 2619 ) ) ) 2620 2625 after after IN 2626 2634 addition addition NN 2635 2637 of of IN 2638 2641 TPA TPA NNP 2642 2644 in in IN 2645 2660 DNA-transfected dna-transfected JJ 2661 2666 U-937 u-937 NN 2667 2672 cells cell NNS 2672 2673 , , , 2674 2684 indicating indicate VBG 2685 2689 that that IN 2690 2693 the the DT 2694 2705 stimulation stimulation NN 2706 2714 appeared appear VBD 2715 2721 likely likely JJ 2722 2724 to to TO 2725 2727 be be VB 2728 2729 a a DT 2730 2734 true true JJ 2735 2742 protein protein NN 2743 2749 kinase kinase NNP 2750 2760 C-mediated c-mediated JJ 2761 2767 signal signal NN 2768 2780 transduction transduction NN 2781 2786 event event NN 2787 2793 rather rather RB 2794 2798 than than IN 2799 2800 a a DT 2801 2816 differentiation differentiation NN 2817 2825 response response NN 2825 2826 . . . 2827 2833 Slight slight JJ 2834 2845 differences difference NNS 2846 2848 in in IN 2849 2852 the the DT 2853 2861 sequence sequence NN 2862 2864 of of IN 2865 2868 the the DT 2869 2873 core core NN 2874 2877 SRF srf NN 2878 2885 binding binding NN 2886 2890 site site NN 2891 2899 compared compare VBN 2900 2904 with with IN 2905 2909 that that DT 2910 2912 of of IN 2913 2916 the the DT 2917 2926 classical classical JJ 2927 2932 c-Fos c-fos NN 2933 2941 promoter promoter NN 2942 2947 serum serum NN 2948 2956 response response NN 2957 2964 element element NN 2964 2965 , , , 2966 2974 together together RB 2975 2979 with with IN 2980 2991 differences difference NNS 2992 2994 in in IN 2995 2998 the the DT 2999 3006 spacing spacing NN 3007 3014 between between IN 3015 3018 the the DT 3019 3022 SRF SRF NNP 3023 3026 and and CC 3027 3030 ETS ETS NNP 3031 3037 motifs motif NNS 3037 3038 , , , 3039 3045 appear appear VBP 3046 3048 to to TO 3049 3056 account account VB 3057 3060 for for IN 3061 3064 the the DT 3065 3074 inability inability NN 3075 3077 of of IN 3078 3081 the the DT 3082 3086 SCMV SCMV NNP 3087 3091 SNEs SNE NNP 3092 3094 to to TO 3095 3102 respond respond VB 3103 3105 to to TO 3106 3111 serum serum NN 3112 3121 induction induction NN 3121 3122 . . .