0	14	Identification	identification	NN
15	17	of	of	IN
18	26	upstream	upstream	JJ
27	37	regulatory	regulatory	JJ
38	46	elements	element	NNS
47	51	that	that	WDT
52	59	repress	repress	VBP
60	70	expression	expression	NN
71	73	of	of	IN
74	79	adult	adult	JJ
80	89	beta-like	beta-like	JJ
90	96	globin	globin	NN
97	102	genes	gene	NNS
103	105	in	in	IN
106	107	a	a	DT
108	117	primitive	primitive	JJ
118	127	erythroid	erythroid	JJ
128	139	environment	environment	NN
139	140	.	.	.

142	145	Our	our	PRP$
146	160	investigations	investigation	NNS
161	165	have	have	VBP
166	173	focused	focus	VBN
174	176	on	on	IN
177	187	localizing	localize	VBG
188	200	cis-elements	cis-element	NNS
201	212	responsible	responsible	JJ
213	216	for	for	IN
217	220	the	the	DT
221	225	down	down	JJ
226	236	regulation	regulation	NN
237	239	of	of	IN
240	243	the	the	DT
244	249	adult	adult	JJ
250	259	beta-like	beta-like	JJ
260	266	globin	globin	NN
267	272	genes	gene	NNS
273	274	(	(	(
274	279	delta	delta	NN
280	283	and	and	CC
284	288	beta	beta	NN
288	289	)	)	)
290	292	in	in	IN
293	301	immature	immature	JJ
301	302	,	,	,
303	305	or	or	CC
306	315	primitive	primitive	JJ
316	325	erythroid	erythroid	JJ
326	333	tissues	tissue	NNS
333	334	.	.	.

335	337	We	we	PRP
338	345	studied	study	VBD
346	351	their	their	PRP$
352	360	activity	activity	NN
361	366	after	after	IN
367	379	transfection	transfection	NN
380	384	into	into	IN
385	389	K562	k562	NN
390	395	cells	cell	NNS
395	396	,	,	,
397	399	an	an	DT
400	415	erythroleukemia	erythroleukemia	NN
416	420	cell	cell	NN
421	425	line	line	NN
426	430	with	with	IN
431	433	an	an	DT
434	449	embryonic-fetal	embryonic-fetal	JJ
450	459	phenotype	phenotype	NN
459	460	.	.	.

461	469	Analyzed	analyze	VBN
470	473	DNA	DNA	NNP
474	483	sequences	sequence	NNS
484	492	included	include	VBD
493	498	delta	delta	SYM
499	502	and	and	CC
503	507	beta	beta	SYM
508	509	5	5	CD
509	510	’	'	SYM
511	519	flanking	flanking	NN
520	527	regions	region	NNS
528	537	extending	extend	VBG
538	542	from	from	IN
543	556	approximately	approximately	RB
557	561	-500	-500	CD
562	564	to	to	TO
565	570	+50bp	+50bp	NN
571	572	(	(	(
572	580	promoter	promoter	NN
581	588	regions	region	NNS
588	589	)	)	)
589	590	,	,	,
591	600	truncated	truncate	VBN
601	606	delta	delta	SYM
607	610	and	and	CC
611	615	beta	beta	SYM
616	617	5	5	CD
617	618	’	'	SYM
619	627	flanking	flanking	NN
628	635	regions	region	NNS
636	645	extending	extend	VBG
646	650	from	from	IN
651	664	approximately	approximately	RB
665	669	-250	-250	CD
670	672	to	to	TO
673	676	+50	+50	CD
677	679	bp	bp	NN
679	680	,	,	,
681	684	and	and	CC
685	693	chimeric	chimeric	JJ
694	702	promoter	promoter	NN
703	716	constructions	construction	NNS
716	717	,	,	,
718	723	which	which	WDT
724	733	consisted	consist	VBD
734	736	of	of	IN
737	738	a	a	DT
739	745	distal	distal	JJ
746	751	delta	delta	NN
752	754	or	or	CC
755	759	beta	beta	NN
760	768	fragment	fragment	NN
769	774	fused	fuse	VBN
775	777	to	to	TO
778	779	a	a	DT
780	788	proximal	proximal	JJ
789	793	beta	beta	NN
794	796	or	or	CC
797	802	delta	delta	NN
803	811	sequence	sequence	NN
811	812	.	.	.

813	815	In	in	IN
816	819	CAT	CAT	NNP
820	828	reporter	reporter	NN
829	842	constructions	construction	NNS
843	845	no	no	DT
846	857	appreciable	appreciable	JJ
858	863	level	level	NN
864	866	of	of	IN
867	870	CAT	CAT	NNP
871	879	activity	activity	NN
880	883	was	be	VBD
884	893	supported	support	VBN
894	896	by	by	IN
897	900	the	the	DT
901	905	beta	beta	NN
906	912	globin	globin	NN
913	921	promoter	promoter	NN
921	922	,	,	,
923	926	and	and	CC
927	931	only	only	RB
932	935	low	low	JJ
936	941	level	level	NN
942	950	activity	activity	NN
951	953	by	by	IN
954	957	the	the	DT
958	963	delta	delta	NN
964	972	promoter	promoter	NN
972	973	.	.	.

974	984	Truncation	Truncation	NNP
985	987	of	of	IN
988	991	the	the	DT
992	996	beta	beta	NN
997	1003	globin	globin	NN
1004	1012	promoter	promoter	NN
1013	1016	led	lead	VBD
1017	1019	to	to	TO
1020	1021	a	a	DT
1022	1025	2-3	2-3	CD
1026	1030	fold	fold	JJ
1031	1039	increase	increase	NN
1040	1042	in	in	IN
1043	1051	promoter	promoter	NN
1052	1060	activity	activity	NN
1060	1061	.	.	.

1062	1064	In	in	IN
1065	1073	contrast	contrast	NN
1073	1074	,	,	,
1075	1083	deletion	deletion	NN
1084	1086	of	of	IN
1087	1090	the	the	DT
1091	1099	upstream	upstream	JJ
1100	1107	portion	portion	NN
1108	1110	of	of	IN
1111	1114	the	the	DT
1115	1120	delta	delta	NN
1121	1129	promoter	promoter	NN
1130	1133	led	lead	VBD
1134	1136	to	to	TO
1137	1138	a	a	DT
1139	1141	10	10	CD
1142	1146	fold	fold	JJ
1147	1155	decrease	decrease	NN
1156	1158	in	in	IN
1159	1169	expression	expression	NN
1169	1170	.	.	.

1171	1179	Coupling	coupling	NN
1180	1182	of	of	IN
1183	1186	the	the	DT
1187	1195	upstream	upstream	JJ
1196	1200	beta	beta	NN
1201	1207	globin	globin	NN
1208	1216	sequence	sequence	NN
1217	1221	from	from	IN
1222	1235	approximately	approximately	RB
1236	1240	-500	-500	CD
1241	1243	to	to	TO
1244	1248	-250	-250	CD
1249	1251	bp	bp	NN
1252	1254	to	to	TO
1255	1258	the	the	DT
1259	1268	truncated	truncate	VBN
1269	1274	delta	delta	NN
1275	1283	promoter	promoter	NN
1284	1292	fragment	fragment	NN
1293	1296	led	lead	VBD
1297	1299	to	to	TO
1300	1308	complete	complete	VB
1309	1319	extinction	extinction	NN
1320	1322	of	of	IN
1323	1336	transcription	transcription	NN
1337	1345	activity	activity	NN
1345	1346	,	,	,
1347	1357	consistent	consistent	JJ
1358	1362	with	with	IN
1363	1364	a	a	DT
1365	1373	negative	negative	JJ
1374	1384	regulatory	regulatory	JJ
1385	1391	effect	effect	NN
1392	1394	of	of	IN
1395	1398	the	the	DT
1399	1403	beta	beta	NN
1404	1410	globin	globin	NN
1411	1415	gene	gene	NN
1416	1424	upstream	upstream	JJ
1425	1435	element(s)	element(s)	NN
1435	1436	.	.	.

1437	1443	Fusion	fusion	NN
1444	1446	of	of	IN
1447	1450	the	the	DT
1451	1459	upstream	upstream	JJ
1460	1467	portion	portion	NN
1468	1470	of	of	IN
1471	1474	the	the	DT
1475	1480	delta	delta	NN
1481	1489	promoter	promoter	NN
1490	1492	to	to	TO
1493	1496	the	the	DT
1497	1506	truncated	truncate	VBN
1507	1511	beta	beta	NN
1512	1518	globin	globin	NN
1519	1527	promoter	promoter	NN
1528	1535	yielded	yield	VBD
1536	1537	a	a	DT
1538	1544	modest	modest	JJ
1545	1553	increase	increase	NN
1554	1556	in	in	IN
1557	1565	promoter	promoter	NN
1566	1574	strength	strength	NN
1575	1583	relative	relative	JJ
1584	1586	to	to	TO
1587	1590	the	the	DT
1591	1600	truncated	truncate	VBN
1601	1605	beta	beta	NN
1606	1610	gene	gene	NN
1611	1619	promoter	promoter	NN
1619	1620	,	,	,
1621	1631	indicating	indicate	VBG
1632	1635	the	the	DT
1636	1644	presence	presence	NN
1645	1647	of	of	IN
1648	1649	a	a	DT
1650	1658	positive	positive	JJ
1659	1674	transcriptional	transcriptional	JJ
1675	1685	element(s)	element(s)	NN
1686	1688	in	in	IN
1689	1692	the	the	DT
1693	1701	upstream	upstream	JJ
1702	1707	delta	delta	NN
1708	1714	globin	globin	NN
1715	1725	regulatory	regulatory	JJ
1726	1732	region	region	NN
1732	1733	.	.	.

1734	1747	Site-directed	site-directed	JJ
1748	1759	mutagenesis	mutagenesis	NN
1760	1762	of	of	IN
1763	1770	binding	bind	VBG
1771	1776	sites	site	NNS
1777	1780	for	for	IN
1781	1784	the	the	DT
1785	1794	repressor	repressor	NN
1795	1803	proteins	protein	NNS
1804	1807	BP1	bp1	NN
1808	1811	and	and	CC
1812	1815	BP2	bp2	NN
1816	1818	in	in	IN
1819	1822	the	the	DT
1823	1831	upstream	upstream	JJ
1832	1839	portion	portion	NN
1840	1842	of	of	IN
1843	1846	the	the	DT
1847	1851	beta	beta	NN
1852	1858	globin	globin	NN
1859	1863	gene	gene	NN
1864	1872	flanking	flanking	NN
1873	1879	region	region	NN
1880	1883	led	lead	VBD
1884	1886	to	to	TO
1887	1888	a	a	DT
1889	1892	4-6	4-6	CD
1893	1897	fold	fold	JJ
1898	1906	increase	increase	NN
1907	1909	in	in	IN
1910	1918	promoter	promoter	NN
1919	1927	activity	activity	NN
1927	1928	.	.	.

1929	1934	DNase	dnase	NN
1935	1936	I	i	CD
1937	1949	footprinting	footprinting	NN
1950	1952	of	of	IN
1953	1956	the	the	DT
1957	1965	upstream	upstream	JJ
1966	1978	delta-globin	delta-globin	NN
1979	1985	region	region	NN
1986	1994	revealed	reveal	VBD
1995	2004	protected	protected	JJ
2005	2014	sequences	sequence	NNS
2015	2028	corresponding	correspond	VBG
2029	2031	to	to	TO
2032	2041	consensus	consensus	NN
2042	2049	binding	binding	NN
2050	2055	sites	site	NNS
2056	2059	for	for	IN
2060	2066	GATA-1	gata-1	NN
2067	2070	and	and	CC
2071	2074	BP2	bp2	NN
2074	2075	.	.	.

2076	2081	These	these	DT
2082	2089	results	result	NNS
2090	2097	confirm	confirm	VBP
2098	2102	that	that	IN
2103	2112	sequences	sequence	NNS
2113	2115	in	in	IN
2116	2119	the	the	DT
2120	2128	upstream	upstream	JJ
2129	2137	promoter	promoter	NN
2138	2144	region	region	NN
2145	2147	of	of	IN
2148	2151	the	the	DT
2152	2157	adult	adult	JJ
2158	2162	beta	beta	NN
2163	2169	globin	globin	NN
2170	2174	gene	gene	NN
2175	2185	contribute	contribute	VBP
2186	2188	to	to	TO
2189	2192	its	its	PRP$
2193	2208	factor-mediated	factor-mediated	JJ
2209	2220	suppression	suppression	NN
2221	2226	early	early	RB
2227	2229	in	in	IN
2230	2241	development	development	NN
2242	2245	and	and	CC
2246	2250	then	then	RB
2251	2254	may	may	MD
2255	2263	modulate	modulate	VB
2264	2267	its	its	PRP$
2268	2278	expression	expression	NN
2279	2281	at	at	IN
2282	2283	a	a	DT
2284	2289	later	later	JJ
2290	2295	stage	stage	NN
2295	2296	.	.	.