PMC:523229 / 40394-50514 JSON TXT 5 Projects

Annnotations TAB TSV DIC JSON TextAE

Id	Subject	Object	Predicate	Lexical cue
T9223	11-19	JJ	denotes	Previous
T9224	20-27	NNS	denotes	studies
T9225	28-35	VBP	denotes	suggest
T9226	36-40	IN	denotes	that
T9228	41-44	NN	denotes	BMP
T9229	45-54	NN	denotes	signaling
T9230	55-57	VBZ	denotes	is
T9227	58-66	VBN	denotes	involved
T9231	67-69	IN	denotes	in
T9232	70-71	DT	denotes	a
T9234	72-77	JJ	denotes	large
T9233	78-84	NN	denotes	number
T9235	85-87	IN	denotes	of
T9236	88-101	JJ	denotes	developmental
T9237	102-108	NNS	denotes	events
T9238	108-109	.	denotes	.
T9239	109-247	sentence	denotes	Many of these events occur early in embryogenesis, and complete inactivation of BMP receptors causes death by E9.5 (Mishina et al. 1995).
T9240	110-114	JJ	denotes	Many
T9242	115-117	IN	denotes	of
T9243	118-123	DT	denotes	these
T9244	124-130	NNS	denotes	events
T9241	131-136	VBP	denotes	occur
T9245	137-142	RB	denotes	early
T9246	143-145	IN	denotes	in
T9247	146-159	NN	denotes	embryogenesis
T9248	159-161	,	denotes	,
T9249	161-164	CC	denotes	and
T9250	165-173	JJ	denotes	complete
T9251	174-186	NN	denotes	inactivation
T9253	187-189	IN	denotes	of
T9254	190-193	NN	denotes	BMP
T9255	194-203	NNS	denotes	receptors
T9252	204-210	VBZ	denotes	causes
T9256	211-216	NN	denotes	death
T9257	217-219	IN	denotes	by
T9258	220-224	NN	denotes	E9.5
T9259	225-226	-LRB-	denotes	(
T9260	226-233	NNP	denotes	Mishina
T9261	234-236	FW	denotes	et
T9262	237-240	FW	denotes	al.
T9263	241-245	CD	denotes	1995
T9264	245-246	-RRB-	denotes	)
T9265	246-247	.	denotes	.
T9266	247-446	sentence	denotes	The Gdf5-Cre recombination system bypasses the early embryonic lethality of Bmpr1a mutations, and provides important new information about the role of this receptor in limb and skeletal development.
T9267	248-251	DT	denotes	The
T9269	252-256	NN	denotes	Gdf5
T9271	256-257	HYPH	denotes	-
T9270	257-260	NN	denotes	Cre
T9272	261-274	NN	denotes	recombination
T9268	275-281	NN	denotes	system
T9273	282-290	VBZ	denotes	bypasses
T9274	291-294	DT	denotes	the
T9276	295-300	JJ	denotes	early
T9277	301-310	JJ	denotes	embryonic
T9275	311-320	NN	denotes	lethality
T9278	321-323	IN	denotes	of
T9279	324-330	NN	denotes	Bmpr1a
T9280	331-340	NNS	denotes	mutations
T9281	340-342	,	denotes	,
T9282	342-345	CC	denotes	and
T9283	346-354	VBZ	denotes	provides
T9284	355-364	JJ	denotes	important
T9286	365-368	JJ	denotes	new
T9285	369-380	NN	denotes	information
T9287	381-386	IN	denotes	about
T9288	387-390	DT	denotes	the
T9289	391-395	NN	denotes	role
T9290	396-398	IN	denotes	of
T9291	399-403	DT	denotes	this
T9292	404-412	NN	denotes	receptor
T9293	413-415	IN	denotes	in
T9294	416-420	NN	denotes	limb
T9296	421-424	CC	denotes	and
T9297	425-433	JJ	denotes	skeletal
T9295	434-445	NN	denotes	development
T9298	445-446	.	denotes	.
T9299	446-710	sentence	denotes	The three major limb phenotypes revealed by eliminating Bmpr1a with Gdf5-driven Cre include webbing between digits, lack of joint formation at specific locations in the ankle, and failure to maintain articular cartilage after birth, resulting in severe arthritis.
T9300	447-450	DT	denotes	The
T9302	451-456	CD	denotes	three
T9303	457-462	JJ	denotes	major
T9304	463-467	NN	denotes	limb
T9301	468-478	NNS	denotes	phenotypes
T9306	479-487	VBN	denotes	revealed
T9307	488-490	IN	denotes	by
T9308	491-502	VBG	denotes	eliminating
T9309	503-509	NN	denotes	Bmpr1a
T9310	510-514	IN	denotes	with
T9311	515-519	NN	denotes	Gdf5
T9313	519-520	HYPH	denotes	-
T9312	520-526	VBN	denotes	driven
T9314	527-530	NN	denotes	Cre
T9305	531-538	VBP	denotes	include
T9315	539-546	NN	denotes	webbing
T9316	547-554	IN	denotes	between
T9317	555-561	NNS	denotes	digits
T9318	561-563	,	denotes	,
T9319	563-567	NN	denotes	lack
T9320	568-570	IN	denotes	of
T9321	571-576	NN	denotes	joint
T9322	577-586	NN	denotes	formation
T9323	587-589	IN	denotes	at
T9324	590-598	JJ	denotes	specific
T9325	599-608	NNS	denotes	locations
T9326	609-611	IN	denotes	in
T9327	612-615	DT	denotes	the
T9328	616-621	NN	denotes	ankle
T9329	621-623	,	denotes	,
T9330	623-626	CC	denotes	and
T9331	627-634	NN	denotes	failure
T9332	635-637	TO	denotes	to
T9333	638-646	VB	denotes	maintain
T9334	647-656	JJ	denotes	articular
T9335	657-666	NN	denotes	cartilage
T9336	667-672	IN	denotes	after
T9337	673-678	NN	denotes	birth
T9338	678-680	,	denotes	,
T9339	680-689	VBG	denotes	resulting
T9340	690-692	IN	denotes	in
T9341	693-699	JJ	denotes	severe
T9342	700-709	NN	denotes	arthritis
T9343	709-710	.	denotes	.
T9344	710-1032	sentence	denotes	Previous studies have shown that manipulation of BMP signaling alters interdigital apoptosis during development of the limb, but no experiment has identified a specific member of the BMP signaling pathway that is required for this process (Yokouchi et al. 1996; Zou and Niswander 1996; Zou et al. 1997; Guha et al. 2002).
T9345	711-719	JJ	denotes	Previous
T9346	720-727	NNS	denotes	studies
T9348	728-732	VBP	denotes	have
T9347	733-738	VBN	denotes	shown
T9349	739-743	IN	denotes	that
T9351	744-756	NN	denotes	manipulation
T9352	757-759	IN	denotes	of
T9353	760-763	NN	denotes	BMP
T9354	764-773	NN	denotes	signaling
T9350	774-780	VBZ	denotes	alters
T9355	781-793	JJ	denotes	interdigital
T9356	794-803	NN	denotes	apoptosis
T9357	804-810	IN	denotes	during
T9358	811-822	NN	denotes	development
T9359	823-825	IN	denotes	of
T9360	826-829	DT	denotes	the
T9361	830-834	NN	denotes	limb
T9362	834-836	,	denotes	,
T9363	836-839	CC	denotes	but
T9364	840-842	DT	denotes	no
T9365	843-853	NN	denotes	experiment
T9367	854-857	VBZ	denotes	has
T9366	858-868	VBN	denotes	identified
T9368	869-870	DT	denotes	a
T9370	871-879	JJ	denotes	specific
T9369	880-886	NN	denotes	member
T9371	887-889	IN	denotes	of
T9372	890-893	DT	denotes	the
T9374	894-897	NN	denotes	BMP
T9375	898-907	NN	denotes	signaling
T9373	908-915	NN	denotes	pathway
T9376	916-920	WDT	denotes	that
T9378	921-923	VBZ	denotes	is
T9377	924-932	VBN	denotes	required
T9379	933-936	IN	denotes	for
T9380	937-941	DT	denotes	this
T9381	942-949	NN	denotes	process
T9382	950-951	-LRB-	denotes	(
T9383	951-959	NNP	denotes	Yokouchi
T9384	960-962	FW	denotes	et
T9385	963-966	FW	denotes	al.
T9386	967-971	CD	denotes	1996
T9387	971-972	:	denotes	;
T9388	973-976	NNP	denotes	Zou
T9389	977-980	CC	denotes	and
T9390	981-990	NNP	denotes	Niswander
T9391	991-995	CD	denotes	1996
T9392	995-996	:	denotes	;
T9393	997-1000	NNP	denotes	Zou
T9394	1001-1003	FW	denotes	et
T9395	1004-1007	FW	denotes	al.
T9396	1008-1012	CD	denotes	1997
T9397	1012-1013	:	denotes	;
T9398	1014-1018	NNP	denotes	Guha
T9399	1019-1021	FW	denotes	et
T9400	1022-1025	FW	denotes	al.
T9401	1026-1030	CD	denotes	2002
T9402	1030-1031	-RRB-	denotes	)
T9403	1031-1032	.	denotes	.
T9404	1032-1204	sentence	denotes	Our new loss-of-function data confirm that BMP signaling is required for interdigital apoptosis and suggests that Bmpr1a is a critical component for mediating this signal.
T9405	1033-1036	PRP$	denotes	Our
T9407	1037-1040	JJ	denotes	new
T9408	1041-1045	NN	denotes	loss
T9409	1045-1046	HYPH	denotes	-
T9410	1046-1048	IN	denotes	of
T9411	1048-1049	HYPH	denotes	-
T9412	1049-1057	NN	denotes	function
T9406	1058-1062	NNS	denotes	data
T9413	1063-1070	VBP	denotes	confirm
T9414	1071-1075	IN	denotes	that
T9416	1076-1079	NN	denotes	BMP
T9417	1080-1089	NN	denotes	signaling
T9418	1090-1092	VBZ	denotes	is
T9415	1093-1101	VBN	denotes	required
T9419	1102-1105	IN	denotes	for
T9420	1106-1118	JJ	denotes	interdigital
T9421	1119-1128	NN	denotes	apoptosis
T9422	1129-1132	CC	denotes	and
T9423	1133-1141	VBZ	denotes	suggests
T9424	1142-1146	IN	denotes	that
T9426	1147-1153	NN	denotes	Bmpr1a
T9425	1154-1156	VBZ	denotes	is
T9427	1157-1158	DT	denotes	a
T9429	1159-1167	JJ	denotes	critical
T9428	1168-1177	NN	denotes	component
T9430	1178-1181	IN	denotes	for
T9431	1182-1191	VBG	denotes	mediating
T9432	1192-1196	DT	denotes	this
T9433	1197-1203	NN	denotes	signal
T9434	1203-1204	.	denotes	.
T9435	1204-1383	sentence	denotes	At some sites, loss of Bmpr1a function leads to a defect in the early stages of joint formation, resulting in a complete failure to form a joint and fusion of bones in the ankle.
T9436	1205-1207	IN	denotes	At
T9438	1208-1212	DT	denotes	some
T9439	1213-1218	NNS	denotes	sites
T9440	1218-1220	,	denotes	,
T9441	1220-1224	NN	denotes	loss
T9442	1225-1227	IN	denotes	of
T9443	1228-1234	NN	denotes	Bmpr1a
T9444	1235-1243	NN	denotes	function
T9437	1244-1249	VBZ	denotes	leads
T9445	1250-1252	IN	denotes	to
T9446	1253-1254	DT	denotes	a
T9447	1255-1261	NN	denotes	defect
T9448	1262-1264	IN	denotes	in
T9449	1265-1268	DT	denotes	the
T9451	1269-1274	JJ	denotes	early
T9450	1275-1281	NNS	denotes	stages
T9452	1282-1284	IN	denotes	of
T9453	1285-1290	NN	denotes	joint
T9454	1291-1300	NN	denotes	formation
T9455	1300-1302	,	denotes	,
T9456	1302-1311	VBG	denotes	resulting
T9457	1312-1314	IN	denotes	in
T9458	1315-1316	DT	denotes	a
T9460	1317-1325	JJ	denotes	complete
T9459	1326-1333	NN	denotes	failure
T9461	1334-1336	TO	denotes	to
T9462	1337-1341	VB	denotes	form
T9463	1342-1343	DT	denotes	a
T9464	1344-1349	NN	denotes	joint
T9465	1350-1353	CC	denotes	and
T9466	1354-1360	NN	denotes	fusion
T9467	1361-1363	IN	denotes	of
T9468	1364-1369	NNS	denotes	bones
T9469	1370-1372	IN	denotes	in
T9470	1373-1376	DT	denotes	the
T9471	1377-1382	NN	denotes	ankle
T9472	1382-1383	.	denotes	.
T9473	1383-1677	sentence	denotes	Mutations in two different ligands in the BMP family, Gdf5 and Gdf6, the Bmpr1b receptor, and in the human Noggin locus (Storm and Kingsley 1996; Gong et al. 1999; Baur et al. 2000; Yi et al. 2000; Settle et al. 2003) also produce defects in joint formation at specific locations in the limbs.
T9474	1384-1393	NNS	denotes	Mutations
T9476	1394-1396	IN	denotes	in
T9477	1397-1400	CD	denotes	two
T9479	1401-1410	JJ	denotes	different
T9478	1411-1418	NNS	denotes	ligands
T9480	1419-1421	IN	denotes	in
T9481	1422-1425	DT	denotes	the
T9483	1426-1429	NN	denotes	BMP
T9482	1430-1436	NN	denotes	family
T9484	1436-1438	,	denotes	,
T9485	1438-1442	NN	denotes	Gdf5
T9486	1443-1446	CC	denotes	and
T9487	1447-1451	NN	denotes	Gdf6
T9488	1451-1453	,	denotes	,
T9489	1453-1456	DT	denotes	the
T9491	1457-1463	NN	denotes	Bmpr1b
T9490	1464-1472	NN	denotes	receptor
T9492	1472-1474	,	denotes	,
T9493	1474-1477	CC	denotes	and
T9494	1478-1480	IN	denotes	in
T9495	1481-1484	DT	denotes	the
T9497	1485-1490	JJ	denotes	human
T9498	1491-1497	NN	denotes	Noggin
T9496	1498-1503	NN	denotes	locus
T9499	1504-1505	-LRB-	denotes	(
T9500	1505-1510	NNP	denotes	Storm
T9501	1511-1514	CC	denotes	and
T9502	1515-1523	NNP	denotes	Kingsley
T9503	1524-1528	CD	denotes	1996
T9504	1528-1529	:	denotes	;
T9505	1530-1534	NNP	denotes	Gong
T9506	1535-1537	FW	denotes	et
T9507	1538-1541	FW	denotes	al.
T9508	1542-1546	CD	denotes	1999
T9509	1546-1547	:	denotes	;
T9510	1548-1552	NNP	denotes	Baur
T9511	1553-1555	FW	denotes	et
T9512	1556-1559	FW	denotes	al.
T9513	1560-1564	CD	denotes	2000
T9514	1564-1565	:	denotes	;
T9515	1566-1568	NNP	denotes	Yi
T9516	1569-1571	FW	denotes	et
T9517	1572-1575	FW	denotes	al.
T9518	1576-1580	CD	denotes	2000
T9519	1580-1581	:	denotes	;
T9520	1582-1588	NNP	denotes	Settle
T9521	1589-1591	FW	denotes	et
T9522	1592-1595	FW	denotes	al.
T9523	1596-1600	CD	denotes	2003
T9524	1600-1601	-RRB-	denotes	)
T9525	1602-1606	RB	denotes	also
T9475	1607-1614	VBP	denotes	produce
T9526	1615-1622	NNS	denotes	defects
T9527	1623-1625	IN	denotes	in
T9528	1626-1631	NN	denotes	joint
T9529	1632-1641	NN	denotes	formation
T9530	1642-1644	IN	denotes	at
T9531	1645-1653	JJ	denotes	specific
T9532	1654-1663	NNS	denotes	locations
T9533	1664-1666	IN	denotes	in
T9534	1667-1670	DT	denotes	the
T9535	1671-1676	NNS	denotes	limbs
T9536	1676-1677	.	denotes	.
T9537	1677-1871	sentence	denotes	The joint defects associated with multiple components of the BMP pathway provide strong evidence that BMP signaling is required for early stages of joint formation at some anatomical locations.
T9538	1678-1681	DT	denotes	The
T9540	1682-1687	NN	denotes	joint
T9539	1688-1695	NNS	denotes	defects
T9542	1696-1706	VBN	denotes	associated
T9543	1707-1711	IN	denotes	with
T9544	1712-1720	JJ	denotes	multiple
T9545	1721-1731	NNS	denotes	components
T9546	1732-1734	IN	denotes	of
T9547	1735-1738	DT	denotes	the
T9549	1739-1742	NN	denotes	BMP
T9548	1743-1750	NN	denotes	pathway
T9541	1751-1758	VBP	denotes	provide
T9550	1759-1765	JJ	denotes	strong
T9551	1766-1774	NN	denotes	evidence
T9552	1775-1779	IN	denotes	that
T9554	1780-1783	NN	denotes	BMP
T9555	1784-1793	NN	denotes	signaling
T9556	1794-1796	VBZ	denotes	is
T9553	1797-1805	VBN	denotes	required
T9557	1806-1809	IN	denotes	for
T9558	1810-1815	JJ	denotes	early
T9559	1816-1822	NNS	denotes	stages
T9560	1823-1825	IN	denotes	of
T9561	1826-1831	NN	denotes	joint
T9562	1832-1841	NN	denotes	formation
T9563	1842-1844	IN	denotes	at
T9564	1845-1849	DT	denotes	some
T9566	1850-1860	JJ	denotes	anatomical
T9565	1861-1870	NNS	denotes	locations
T9567	1870-1871	.	denotes	.
T9568	1871-1958	sentence	denotes	Most joints still form normally when Bmpr1a is knocked out in Gdf5 expression domains.
T9569	1872-1876	JJS	denotes	Most
T9570	1877-1883	NNS	denotes	joints
T9572	1884-1889	RB	denotes	still
T9571	1890-1894	VBP	denotes	form
T9573	1895-1903	RB	denotes	normally
T9574	1904-1908	WRB	denotes	when
T9576	1909-1915	NN	denotes	Bmpr1a
T9577	1916-1918	VBZ	denotes	is
T9575	1919-1926	VBN	denotes	knocked
T9578	1927-1930	RP	denotes	out
T9579	1931-1933	IN	denotes	in
T9580	1934-1938	NN	denotes	Gdf5
T9582	1939-1949	NN	denotes	expression
T9581	1950-1957	NNS	denotes	domains
T9583	1957-1958	.	denotes	.
T9584	1958-2278	sentence	denotes	The lack of joint fusions outside the ankle region could be due to differences in requirement for BMP signaling in different joints, to compensating expression of other BMP receptors outside the ankles, or to differences in the detailed timing of Gdf5-Cre stimulated gene inactivation in ankles and other joint regions.
T9585	1959-1962	DT	denotes	The
T9586	1963-1967	NN	denotes	lack
T9588	1968-1970	IN	denotes	of
T9589	1971-1976	NN	denotes	joint
T9590	1977-1984	NNS	denotes	fusions
T9591	1985-1992	IN	denotes	outside
T9592	1993-1996	DT	denotes	the
T9594	1997-2002	NN	denotes	ankle
T9593	2003-2009	NN	denotes	region
T9595	2010-2015	MD	denotes	could
T9587	2016-2018	VB	denotes	be
T9596	2019-2022	IN	denotes	due
T9597	2023-2025	IN	denotes	to
T9598	2026-2037	NNS	denotes	differences
T9599	2038-2040	IN	denotes	in
T9600	2041-2052	NN	denotes	requirement
T9601	2053-2056	IN	denotes	for
T9602	2057-2060	NN	denotes	BMP
T9603	2061-2070	NN	denotes	signaling
T9604	2071-2073	IN	denotes	in
T9605	2074-2083	JJ	denotes	different
T9606	2084-2090	NNS	denotes	joints
T9607	2090-2092	,	denotes	,
T9608	2092-2094	IN	denotes	to
T9609	2095-2107	VBG	denotes	compensating
T9610	2108-2118	NN	denotes	expression
T9611	2119-2121	IN	denotes	of
T9612	2122-2127	JJ	denotes	other
T9614	2128-2131	NN	denotes	BMP
T9613	2132-2141	NNS	denotes	receptors
T9615	2142-2149	IN	denotes	outside
T9616	2150-2153	DT	denotes	the
T9617	2154-2160	NNS	denotes	ankles
T9618	2160-2162	,	denotes	,
T9619	2162-2164	CC	denotes	or
T9620	2165-2167	IN	denotes	to
T9621	2168-2179	NNS	denotes	differences
T9622	2180-2182	IN	denotes	in
T9623	2183-2186	DT	denotes	the
T9625	2187-2195	VBN	denotes	detailed
T9624	2196-2202	NN	denotes	timing
T9626	2203-2205	IN	denotes	of
T9627	2206-2210	NN	denotes	Gdf5
T9629	2210-2211	HYPH	denotes	-
T9628	2211-2214	NN	denotes	Cre
T9630	2215-2225	VBN	denotes	stimulated
T9632	2226-2230	NN	denotes	gene
T9631	2231-2243	NN	denotes	inactivation
T9633	2244-2246	IN	denotes	in
T9634	2247-2253	NNS	denotes	ankles
T9635	2254-2257	CC	denotes	and
T9636	2258-2263	JJ	denotes	other
T9638	2264-2269	NN	denotes	joint
T9637	2270-2277	NNS	denotes	regions
T9639	2277-2278	.	denotes	.
T9640	2278-2572	sentence	denotes	Comparison of the expression of the HPLAP marker (driven directly by Gdf5 control elements) and the R26R LACZ marker (expressed following Gdf5-Cre recombination) suggests that recombination-stimulated changes in gene expression may be delayed for a 0.5–1 d in the digit region (see Figure 1C).
T9641	2279-2289	NN	denotes	Comparison
T9643	2290-2292	IN	denotes	of
T9644	2293-2296	DT	denotes	the
T9645	2297-2307	NN	denotes	expression
T9646	2308-2310	IN	denotes	of
T9647	2311-2314	DT	denotes	the
T9649	2315-2320	NN	denotes	HPLAP
T9648	2321-2327	NN	denotes	marker
T9650	2328-2329	-LRB-	denotes	(
T9651	2329-2335	VBN	denotes	driven
T9652	2336-2344	RB	denotes	directly
T9653	2345-2347	IN	denotes	by
T9654	2348-2352	NN	denotes	Gdf5
T9656	2353-2360	NN	denotes	control
T9655	2361-2369	NNS	denotes	elements
T9657	2369-2370	-RRB-	denotes	)
T9658	2371-2374	CC	denotes	and
T9659	2375-2378	DT	denotes	the
T9661	2379-2383	NN	denotes	R26R
T9662	2384-2388	NN	denotes	LACZ
T9660	2389-2395	NN	denotes	marker
T9663	2396-2397	-LRB-	denotes	(
T9664	2397-2406	VBN	denotes	expressed
T9665	2407-2416	VBG	denotes	following
T9666	2417-2421	NN	denotes	Gdf5
T9668	2421-2422	HYPH	denotes	-
T9667	2422-2425	NN	denotes	Cre
T9669	2426-2439	NN	denotes	recombination
T9670	2439-2440	-RRB-	denotes	)
T9642	2441-2449	VBZ	denotes	suggests
T9671	2450-2454	IN	denotes	that
T9673	2455-2468	NN	denotes	recombination
T9675	2468-2469	HYPH	denotes	-
T9674	2469-2479	VBN	denotes	stimulated
T9676	2480-2487	NNS	denotes	changes
T9677	2488-2490	IN	denotes	in
T9678	2491-2495	NN	denotes	gene
T9679	2496-2506	NN	denotes	expression
T9680	2507-2510	MD	denotes	may
T9681	2511-2513	VB	denotes	be
T9672	2514-2521	VBN	denotes	delayed
T9682	2522-2525	IN	denotes	for
T9683	2526-2527	DT	denotes	a
T9685	2528-2531	CD	denotes	0.5
T9687	2531-2532	SYM	denotes	–
T9686	2532-2533	CD	denotes	1
T9684	2534-2535	NN	denotes	d
T9688	2536-2538	IN	denotes	in
T9689	2539-2542	DT	denotes	the
T9691	2543-2548	NN	denotes	digit
T9690	2549-2555	NN	denotes	region
T9692	2556-2557	-LRB-	denotes	(
T9693	2557-2560	VB	denotes	see
T9694	2561-2567	NN	denotes	Figure
T9695	2568-2570	CD	denotes	1C
T9696	2570-2571	-RRB-	denotes	)
T9697	2571-2572	.	denotes	.
T9698	2572-2789	sentence	denotes	In addition, levels of Bmpr1a mRNA and protein may persist for some time following Gdf5-Cre stimulated recombination, making it possible to bypass an early requirement for Bmpr1a in joint formation at some locations.
T9699	2573-2575	IN	denotes	In
T9701	2576-2584	NN	denotes	addition
T9702	2584-2586	,	denotes	,
T9703	2586-2592	NNS	denotes	levels
T9704	2593-2595	IN	denotes	of
T9705	2596-2602	NN	denotes	Bmpr1a
T9706	2603-2607	NN	denotes	mRNA
T9707	2608-2611	CC	denotes	and
T9708	2612-2619	NN	denotes	protein
T9709	2620-2623	MD	denotes	may
T9700	2624-2631	VB	denotes	persist
T9710	2632-2635	IN	denotes	for
T9711	2636-2640	DT	denotes	some
T9712	2641-2645	NN	denotes	time
T9713	2646-2655	VBG	denotes	following
T9714	2656-2660	NN	denotes	Gdf5
T9716	2660-2661	HYPH	denotes	-
T9715	2661-2664	NN	denotes	Cre
T9717	2665-2675	VBN	denotes	stimulated
T9718	2676-2689	NN	denotes	recombination
T9719	2689-2691	,	denotes	,
T9720	2691-2697	VBG	denotes	making
T9721	2698-2700	PRP	denotes	it
T9722	2701-2709	JJ	denotes	possible
T9723	2710-2712	TO	denotes	to
T9724	2713-2719	VB	denotes	bypass
T9725	2720-2722	DT	denotes	an
T9727	2723-2728	JJ	denotes	early
T9726	2729-2740	NN	denotes	requirement
T9728	2741-2744	IN	denotes	for
T9729	2745-2751	NN	denotes	Bmpr1a
T9730	2752-2754	IN	denotes	in
T9731	2755-2760	NN	denotes	joint
T9732	2761-2770	NN	denotes	formation
T9733	2771-2773	IN	denotes	at
T9734	2774-2778	DT	denotes	some
T9735	2779-2788	NNS	denotes	locations
T9736	2788-2789	.	denotes	.
T9737	2789-2939	sentence	denotes	Following the decay of Bmpr1a mRNA and protein, the Gdf5-Cre strategy should result in permanent inactivation of Bmpr1a function in recombined cells.
T9738	2790-2799	VBG	denotes	Following
T9740	2800-2803	DT	denotes	the
T9741	2804-2809	NN	denotes	decay
T9742	2810-2812	IN	denotes	of
T9743	2813-2819	NN	denotes	Bmpr1a
T9744	2820-2824	NN	denotes	mRNA
T9745	2825-2828	CC	denotes	and
T9746	2829-2836	NN	denotes	protein
T9747	2836-2838	,	denotes	,
T9748	2838-2841	DT	denotes	the
T9750	2842-2846	NN	denotes	Gdf5
T9752	2846-2847	HYPH	denotes	-
T9751	2847-2850	NN	denotes	Cre
T9749	2851-2859	NN	denotes	strategy
T9753	2860-2866	MD	denotes	should
T9739	2867-2873	VB	denotes	result
T9754	2874-2876	IN	denotes	in
T9755	2877-2886	JJ	denotes	permanent
T9756	2887-2899	NN	denotes	inactivation
T9757	2900-2902	IN	denotes	of
T9758	2903-2909	NN	denotes	Bmpr1a
T9759	2910-2918	NN	denotes	function
T9760	2919-2921	IN	denotes	in
T9761	2922-2932	VBN	denotes	recombined
T9762	2933-2938	NNS	denotes	cells
T9763	2938-2939	.	denotes	.
T9764	2939-3060	sentence	denotes	This system thus provides one of the first strong genetic tests of Bmpr1a function at later stages of joint development.
T9765	2940-2944	DT	denotes	This
T9766	2945-2951	NN	denotes	system
T9768	2952-2956	RB	denotes	thus
T9767	2957-2965	VBZ	denotes	provides
T9769	2966-2969	CD	denotes	one
T9770	2970-2972	IN	denotes	of
T9771	2973-2976	DT	denotes	the
T9773	2977-2982	JJ	denotes	first
T9774	2983-2989	JJ	denotes	strong
T9775	2990-2997	JJ	denotes	genetic
T9772	2998-3003	NNS	denotes	tests
T9776	3004-3006	IN	denotes	of
T9777	3007-3013	NN	denotes	Bmpr1a
T9778	3014-3022	NN	denotes	function
T9779	3023-3025	IN	denotes	at
T9780	3026-3031	JJ	denotes	later
T9781	3032-3038	NNS	denotes	stages
T9782	3039-3041	IN	denotes	of
T9783	3042-3047	NN	denotes	joint
T9784	3048-3059	NN	denotes	development
T9785	3059-3060	.	denotes	.
T9786	3060-3296	sentence	denotes	Despite the normal appearance of articular regions and gene expression immediately after birth, Bmpr1a-deficient animals are unable to maintain the normal differentiated state of articular cartilage as they continue to develop and age.
T9787	3061-3068	IN	denotes	Despite
T9789	3069-3072	DT	denotes	the
T9791	3073-3079	JJ	denotes	normal
T9790	3080-3090	NN	denotes	appearance
T9792	3091-3093	IN	denotes	of
T9793	3094-3103	JJ	denotes	articular
T9794	3104-3111	NNS	denotes	regions
T9795	3112-3115	CC	denotes	and
T9796	3116-3120	NN	denotes	gene
T9797	3121-3131	NN	denotes	expression
T9798	3132-3143	RB	denotes	immediately
T9799	3144-3149	IN	denotes	after
T9800	3150-3155	NN	denotes	birth
T9801	3155-3157	,	denotes	,
T9802	3157-3163	NN	denotes	Bmpr1a
T9804	3163-3164	HYPH	denotes	-
T9803	3164-3173	JJ	denotes	deficient
T9805	3174-3181	NNS	denotes	animals
T9788	3182-3185	VBP	denotes	are
T9806	3186-3192	JJ	denotes	unable
T9807	3193-3195	TO	denotes	to
T9808	3196-3204	VB	denotes	maintain
T9809	3205-3208	DT	denotes	the
T9811	3209-3215	JJ	denotes	normal
T9812	3216-3230	VBN	denotes	differentiated
T9810	3231-3236	NN	denotes	state
T9813	3237-3239	IN	denotes	of
T9814	3240-3249	JJ	denotes	articular
T9815	3250-3259	NN	denotes	cartilage
T9816	3260-3262	IN	denotes	as
T9818	3263-3267	PRP	denotes	they
T9817	3268-3276	VBP	denotes	continue
T9819	3277-3279	TO	denotes	to
T9820	3280-3287	VB	denotes	develop
T9821	3288-3291	CC	denotes	and
T9822	3292-3295	VB	denotes	age
T9823	3295-3296	.	denotes	.
T9824	3296-3442	sentence	denotes	These results suggest that BMP receptor signaling is essential for continued health and integrity of articular cartilage in the postnatal period.
T9825	3297-3302	DT	denotes	These
T9826	3303-3310	NNS	denotes	results
T9827	3311-3318	VBP	denotes	suggest
T9828	3319-3323	IN	denotes	that
T9830	3324-3327	NN	denotes	BMP
T9831	3328-3336	NN	denotes	receptor
T9832	3337-3346	NN	denotes	signaling
T9829	3347-3349	VBZ	denotes	is
T9833	3350-3359	JJ	denotes	essential
T9834	3360-3363	IN	denotes	for
T9835	3364-3373	VBN	denotes	continued
T9836	3374-3380	NN	denotes	health
T9837	3381-3384	CC	denotes	and
T9838	3385-3394	NN	denotes	integrity
T9839	3395-3397	IN	denotes	of
T9840	3398-3407	JJ	denotes	articular
T9841	3408-3417	NN	denotes	cartilage
T9842	3418-3420	IN	denotes	in
T9843	3421-3424	DT	denotes	the
T9845	3425-3434	JJ	denotes	postnatal
T9844	3435-3441	NN	denotes	period
T9846	3441-3442	.	denotes	.
T9847	3442-3592	sentence	denotes	Articular cartilage is a key component of synovial joints and is one of the few regions in the skeleton where cartilage is maintained into adulthood.
T9848	3443-3452	JJ	denotes	Articular
T9849	3453-3462	NN	denotes	cartilage
T9850	3463-3465	VBZ	denotes	is
T9851	3466-3467	DT	denotes	a
T9853	3468-3471	JJ	denotes	key
T9852	3472-3481	NN	denotes	component
T9854	3482-3484	IN	denotes	of
T9855	3485-3493	JJ	denotes	synovial
T9856	3494-3500	NNS	denotes	joints
T9857	3501-3504	CC	denotes	and
T9858	3505-3507	VBZ	denotes	is
T9859	3508-3511	CD	denotes	one
T9860	3512-3514	IN	denotes	of
T9861	3515-3518	DT	denotes	the
T9863	3519-3522	JJ	denotes	few
T9862	3523-3530	NNS	denotes	regions
T9864	3531-3533	IN	denotes	in
T9865	3534-3537	DT	denotes	the
T9866	3538-3546	NN	denotes	skeleton
T9867	3547-3552	WRB	denotes	where
T9869	3553-3562	NN	denotes	cartilage
T9870	3563-3565	VBZ	denotes	is
T9868	3566-3576	VBN	denotes	maintained
T9871	3577-3581	IN	denotes	into
T9872	3582-3591	NN	denotes	adulthood
T9873	3591-3592	.	denotes	.
T9874	3592-3772	sentence	denotes	Despite the importance of articular cartilage in joint health and mobility, little is known about the factors that create and maintain it in thin layers at the ends of long bones.
T9875	3593-3600	IN	denotes	Despite
T9877	3601-3604	DT	denotes	the
T9878	3605-3615	NN	denotes	importance
T9879	3616-3618	IN	denotes	of
T9880	3619-3628	JJ	denotes	articular
T9881	3629-3638	NN	denotes	cartilage
T9882	3639-3641	IN	denotes	in
T9883	3642-3647	NN	denotes	joint
T9884	3648-3654	NN	denotes	health
T9885	3655-3658	CC	denotes	and
T9886	3659-3667	NN	denotes	mobility
T9887	3667-3669	,	denotes	,
T9888	3669-3675	JJ	denotes	little
T9889	3676-3678	VBZ	denotes	is
T9876	3679-3684	VBN	denotes	known
T9890	3685-3690	IN	denotes	about
T9891	3691-3694	DT	denotes	the
T9892	3695-3702	NNS	denotes	factors
T9893	3703-3707	WDT	denotes	that
T9894	3708-3714	VBP	denotes	create
T9895	3715-3718	CC	denotes	and
T9896	3719-3727	VBP	denotes	maintain
T9897	3728-3730	PRP	denotes	it
T9898	3731-3733	IN	denotes	in
T9899	3734-3738	JJ	denotes	thin
T9900	3739-3745	NNS	denotes	layers
T9901	3746-3748	IN	denotes	at
T9902	3749-3752	DT	denotes	the
T9903	3753-3757	NNS	denotes	ends
T9904	3758-3760	IN	denotes	of
T9905	3761-3765	JJ	denotes	long
T9906	3766-3771	NNS	denotes	bones
T9907	3771-3772	.	denotes	.
T9908	3772-4069	sentence	denotes	In our experiments, articular cartilage lacking Bmpr1a retains some normal characteristics, in that it maintains a very low proliferation rate, does not express Col1a1, and continues to express SOX9, a major transcription factor regulating expression of structural components of cartilage matrix.
T9909	3773-3775	IN	denotes	In
T9911	3776-3779	PRP$	denotes	our
T9912	3780-3791	NNS	denotes	experiments
T9913	3791-3793	,	denotes	,
T9914	3793-3802	JJ	denotes	articular
T9915	3803-3812	NN	denotes	cartilage
T9916	3813-3820	VBG	denotes	lacking
T9917	3821-3827	NN	denotes	Bmpr1a
T9910	3828-3835	VBZ	denotes	retains
T9918	3836-3840	DT	denotes	some
T9920	3841-3847	JJ	denotes	normal
T9919	3848-3863	NNS	denotes	characteristics
T9921	3863-3865	,	denotes	,
T9922	3865-3867	IN	denotes	in
T9924	3868-3872	IN	denotes	that
T9925	3873-3875	PRP	denotes	it
T9923	3876-3885	VBZ	denotes	maintains
T9926	3886-3887	DT	denotes	a
T9928	3888-3892	RB	denotes	very
T9929	3893-3896	JJ	denotes	low
T9930	3897-3910	NN	denotes	proliferation
T9927	3911-3915	NN	denotes	rate
T9931	3915-3917	,	denotes	,
T9932	3917-3921	VBZ	denotes	does
T9934	3922-3925	RB	denotes	not
T9933	3926-3933	VB	denotes	express
T9935	3934-3940	NN	denotes	Col1a1
T9936	3940-3942	,	denotes	,
T9937	3942-3945	CC	denotes	and
T9938	3946-3955	VBZ	denotes	continues
T9939	3956-3958	TO	denotes	to
T9940	3959-3966	VB	denotes	express
T9941	3967-3971	NN	denotes	SOX9
T9942	3971-3973	,	denotes	,
T9943	3973-3974	DT	denotes	a
T9945	3975-3980	JJ	denotes	major
T9946	3981-3994	NN	denotes	transcription
T9944	3995-4001	NN	denotes	factor
T9947	4002-4012	VBG	denotes	regulating
T9948	4013-4023	NN	denotes	expression
T9949	4024-4026	IN	denotes	of
T9950	4027-4037	JJ	denotes	structural
T9951	4038-4048	NNS	denotes	components
T9952	4049-4051	IN	denotes	of
T9953	4052-4061	NN	denotes	cartilage
T9954	4062-4068	NN	denotes	matrix
T9955	4068-4069	.	denotes	.
T9956	4069-4259	sentence	denotes	However, several of the most prominent structural components of cartilage matrix fail to be maintained in mutant animals, resulting in decreased synthesis of Col2a1, Agg, and proteoglycans.
T9957	4070-4077	RB	denotes	However
T9959	4077-4079	,	denotes	,
T9960	4079-4086	JJ	denotes	several
T9961	4087-4089	IN	denotes	of
T9962	4090-4093	DT	denotes	the
T9964	4094-4098	RBS	denotes	most
T9965	4099-4108	JJ	denotes	prominent
T9966	4109-4119	JJ	denotes	structural
T9963	4120-4130	NNS	denotes	components
T9967	4131-4133	IN	denotes	of
T9968	4134-4143	NN	denotes	cartilage
T9969	4144-4150	NN	denotes	matrix
T9958	4151-4155	VBP	denotes	fail
T9970	4156-4158	TO	denotes	to
T9972	4159-4161	VB	denotes	be
T9971	4162-4172	VBN	denotes	maintained
T9973	4173-4175	IN	denotes	in
T9974	4176-4182	NN	denotes	mutant
T9975	4183-4190	NNS	denotes	animals
T9976	4190-4192	,	denotes	,
T9977	4192-4201	VBG	denotes	resulting
T9978	4202-4204	IN	denotes	in
T9979	4205-4214	VBN	denotes	decreased
T9980	4215-4224	NN	denotes	synthesis
T9981	4225-4227	IN	denotes	of
T9982	4228-4234	NN	denotes	Col2a1
T9983	4234-4236	,	denotes	,
T9984	4236-4239	NN	denotes	Agg
T9985	4239-4241	,	denotes	,
T9986	4241-4244	CC	denotes	and
T9987	4245-4258	NNS	denotes	proteoglycans
T9988	4258-4259	.	denotes	.
T9989	4259-4378	sentence	denotes	Therefore, BMPR1A appears to maintain articular cartilage primarily through inducing expression of key ECM components.
T9990	4260-4269	RB	denotes	Therefore
T9992	4269-4271	,	denotes	,
T9993	4271-4277	NN	denotes	BMPR1A
T9991	4278-4285	VBZ	denotes	appears
T9994	4286-4288	TO	denotes	to
T9995	4289-4297	VB	denotes	maintain
T9996	4298-4307	JJ	denotes	articular
T9997	4308-4317	NN	denotes	cartilage
T9998	4318-4327	RB	denotes	primarily
T9999	4328-4335	IN	denotes	through
T10000	4336-4344	VBG	denotes	inducing
T10001	4345-4355	NN	denotes	expression
T10002	4356-4358	IN	denotes	of
T10003	4359-4362	JJ	denotes	key
T10005	4363-4366	NN	denotes	ECM
T10004	4367-4377	NNS	denotes	components
T10006	4377-4378	.	denotes	.
T10007	4378-4588	sentence	denotes	It is interesting that the SOX9 transcription factor continues to be expressed in mutant cartilage despite loss of Col2a1, a direct target of this transcription factor (Bell et al. 1997; Lefebvre et al. 1997).
T10008	4379-4381	PRP	denotes	It
T10009	4382-4384	VBZ	denotes	is
T10010	4385-4396	JJ	denotes	interesting
T10011	4397-4401	IN	denotes	that
T10013	4402-4405	DT	denotes	the
T10015	4406-4410	NN	denotes	SOX9
T10016	4411-4424	NN	denotes	transcription
T10014	4425-4431	NN	denotes	factor
T10012	4432-4441	VBZ	denotes	continues
T10017	4442-4444	TO	denotes	to
T10019	4445-4447	VB	denotes	be
T10018	4448-4457	VBN	denotes	expressed
T10020	4458-4460	IN	denotes	in
T10021	4461-4467	NN	denotes	mutant
T10022	4468-4477	NN	denotes	cartilage
T10023	4478-4485	IN	denotes	despite
T10024	4486-4490	NN	denotes	loss
T10025	4491-4493	IN	denotes	of
T10026	4494-4500	NN	denotes	Col2a1
T10027	4500-4502	,	denotes	,
T10028	4502-4503	DT	denotes	a
T10030	4504-4510	JJ	denotes	direct
T10029	4511-4517	NN	denotes	target
T10031	4518-4520	IN	denotes	of
T10032	4521-4525	DT	denotes	this
T10034	4526-4539	NN	denotes	transcription
T10033	4540-4546	NN	denotes	factor
T10035	4547-4548	-LRB-	denotes	(
T10036	4548-4552	NNP	denotes	Bell
T10037	4553-4555	FW	denotes	et
T10038	4556-4559	FW	denotes	al.
T10039	4560-4564	CD	denotes	1997
T10040	4564-4565	:	denotes	;
T10041	4566-4574	NNP	denotes	Lefebvre
T10042	4575-4577	FW	denotes	et
T10043	4578-4581	FW	denotes	al.
T10044	4582-4586	CD	denotes	1997
T10045	4586-4587	-RRB-	denotes	)
T10046	4587-4588	.	denotes	.
T10047	4588-4812	sentence	denotes	Previous studies suggest that SOX9 activity can be modified by protein kinase A (PKA)-dependent protein phosphorylation, or by coexpression of two related proteins, L-SOX5 and SOX6 (Lefebvre et al. 1998; Huang et al. 2000).
T10048	4589-4597	JJ	denotes	Previous
T10049	4598-4605	NNS	denotes	studies
T10050	4606-4613	VBP	denotes	suggest
T10051	4614-4618	IN	denotes	that
T10053	4619-4623	NN	denotes	SOX9
T10054	4624-4632	NN	denotes	activity
T10055	4633-4636	MD	denotes	can
T10056	4637-4639	VB	denotes	be
T10052	4640-4648	VBN	denotes	modified
T10057	4649-4651	IN	denotes	by
T10058	4652-4659	NN	denotes	protein
T10060	4660-4666	NN	denotes	kinase
T10059	4667-4668	NN	denotes	A
T10062	4669-4670	-LRB-	denotes	(
T10063	4670-4673	NN	denotes	PKA
T10064	4673-4674	-RRB-	denotes	)
T10065	4674-4675	HYPH	denotes	-
T10061	4675-4684	JJ	denotes	dependent
T10067	4685-4692	NN	denotes	protein
T10066	4693-4708	NN	denotes	phosphorylation
T10068	4708-4710	,	denotes	,
T10069	4710-4712	CC	denotes	or
T10070	4713-4715	IN	denotes	by
T10071	4716-4728	NN	denotes	coexpression
T10072	4729-4731	IN	denotes	of
T10073	4732-4735	CD	denotes	two
T10075	4736-4743	VBN	denotes	related
T10074	4744-4752	NN	denotes	proteins
T10076	4752-4754	,	denotes	,
T10077	4754-4755	NN	denotes	L
T10079	4755-4756	HYPH	denotes	-
T10078	4756-4760	NN	denotes	SOX5
T10080	4761-4764	CC	denotes	and
T10081	4765-4769	NN	denotes	SOX6
T10082	4770-4771	-LRB-	denotes	(
T10083	4771-4779	NNP	denotes	Lefebvre
T10084	4780-4782	FW	denotes	et
T10085	4783-4786	FW	denotes	al.
T10086	4787-4791	CD	denotes	1998
T10087	4791-4792	:	denotes	;
T10088	4793-4798	NNP	denotes	Huang
T10089	4799-4801	FW	denotes	et
T10090	4802-4805	FW	denotes	al.
T10091	4806-4810	CD	denotes	2000
T10092	4810-4811	-RRB-	denotes	)
T10093	4811-4812	.	denotes	.
T10094	4812-5075	sentence	denotes	In addition, close examination of the order of genes induced during chicken digit formation reveals that Sox9 turns on first, followed by Bmpr1b with L-Sox5, and then Sox6 and the cartilage matrix structural components Col2a1 and Agg (Chimal-Monroy et al. 2003).
T10095	4813-4815	IN	denotes	In
T10097	4816-4824	NN	denotes	addition
T10098	4824-4826	,	denotes	,
T10099	4826-4831	JJ	denotes	close
T10100	4832-4843	NN	denotes	examination
T10101	4844-4846	IN	denotes	of
T10102	4847-4850	DT	denotes	the
T10103	4851-4856	NN	denotes	order
T10104	4857-4859	IN	denotes	of
T10105	4860-4865	NNS	denotes	genes
T10106	4866-4873	VBN	denotes	induced
T10107	4874-4880	IN	denotes	during
T10108	4881-4888	NN	denotes	chicken
T10110	4889-4894	NN	denotes	digit
T10109	4895-4904	NN	denotes	formation
T10096	4905-4912	VBZ	denotes	reveals
T10111	4913-4917	IN	denotes	that
T10113	4918-4922	NN	denotes	Sox9
T10112	4923-4928	VBZ	denotes	turns
T10114	4929-4931	RP	denotes	on
T10115	4932-4937	RB	denotes	first
T10116	4937-4939	,	denotes	,
T10117	4939-4947	VBN	denotes	followed
T10118	4948-4950	IN	denotes	by
T10119	4951-4957	NN	denotes	Bmpr1b
T10120	4958-4962	IN	denotes	with
T10121	4963-4964	NN	denotes	L
T10123	4964-4965	HYPH	denotes	-
T10122	4965-4969	NN	denotes	Sox5
T10124	4969-4971	,	denotes	,
T10125	4971-4974	CC	denotes	and
T10126	4975-4979	RB	denotes	then
T10127	4980-4984	NN	denotes	Sox6
T10128	4985-4988	CC	denotes	and
T10129	4989-4992	DT	denotes	the
T10131	4993-5002	NN	denotes	cartilage
T10132	5003-5009	NN	denotes	matrix
T10133	5010-5020	JJ	denotes	structural
T10130	5021-5031	NNS	denotes	components
T10134	5032-5038	NN	denotes	Col2a1
T10135	5039-5042	CC	denotes	and
T10136	5043-5046	NN	denotes	Agg
T10137	5047-5048	-LRB-	denotes	(
T10138	5048-5054	NNP	denotes	Chimal
T10139	5054-5055	HYPH	denotes	-
T10140	5055-5061	NNP	denotes	Monroy
T10141	5062-5064	FW	denotes	et
T10142	5065-5068	FW	denotes	al.
T10143	5069-5073	CD	denotes	2003
T10144	5073-5074	-RRB-	denotes	)
T10145	5074-5075	.	denotes	.
T10146	5075-5370	sentence	denotes	These results, together with the altered pattern of gene expression seen in our Bmpr1a-deficient mice, suggest that BMPR1A signaling may normally act to stimulate SOX9 by post-translational protein modification, or to induce L-Sox5 or Sox6 in cartilage to maintain expression of ECM components.
T10147	5076-5081	DT	denotes	These
T10148	5082-5089	NNS	denotes	results
T10150	5089-5091	,	denotes	,
T10151	5091-5099	RB	denotes	together
T10152	5100-5104	IN	denotes	with
T10153	5105-5108	DT	denotes	the
T10155	5109-5116	VBN	denotes	altered
T10154	5117-5124	NN	denotes	pattern
T10156	5125-5127	IN	denotes	of
T10157	5128-5132	NN	denotes	gene
T10158	5133-5143	NN	denotes	expression
T10159	5144-5148	VBN	denotes	seen
T10160	5149-5151	IN	denotes	in
T10161	5152-5155	PRP$	denotes	our
T10163	5156-5162	NN	denotes	Bmpr1a
T10165	5162-5163	HYPH	denotes	-
T10164	5163-5172	JJ	denotes	deficient
T10162	5173-5177	NNS	denotes	mice
T10166	5177-5179	,	denotes	,
T10149	5179-5186	VBP	denotes	suggest
T10167	5187-5191	IN	denotes	that
T10169	5192-5198	NN	denotes	BMPR1A
T10170	5199-5208	NN	denotes	signaling
T10171	5209-5212	MD	denotes	may
T10172	5213-5221	RB	denotes	normally
T10168	5222-5225	VB	denotes	act
T10173	5226-5228	TO	denotes	to
T10174	5229-5238	VB	denotes	stimulate
T10175	5239-5243	NN	denotes	SOX9
T10176	5244-5246	IN	denotes	by
T10177	5247-5265	JJ	denotes	post-translational
T10179	5266-5273	NN	denotes	protein
T10178	5274-5286	NN	denotes	modification
T10180	5286-5288	,	denotes	,
T10181	5288-5290	CC	denotes	or
T10182	5291-5293	TO	denotes	to
T10183	5294-5300	VB	denotes	induce
T10184	5301-5302	NN	denotes	L
T10186	5302-5303	HYPH	denotes	-
T10185	5303-5307	NN	denotes	Sox5
T10187	5308-5310	CC	denotes	or
T10188	5311-5315	NN	denotes	Sox6
T10189	5316-5318	IN	denotes	in
T10190	5319-5328	NN	denotes	cartilage
T10191	5329-5331	TO	denotes	to
T10192	5332-5340	VB	denotes	maintain
T10193	5341-5351	NN	denotes	expression
T10194	5352-5354	IN	denotes	of
T10195	5355-5358	NN	denotes	ECM
T10196	5359-5369	NNS	denotes	components
T10197	5369-5370	.	denotes	.
T10198	5370-5560	sentence	denotes	These models are consistent with the ability of BMP2 to both increase PKA activity and induce expression of Sox6 in tissue culture cells (Lee and Chuong 1997; Fernandez-Lloris et al. 2003).
T10199	5371-5376	DT	denotes	These
T10200	5377-5383	NNS	denotes	models
T10201	5384-5387	VBP	denotes	are
T10202	5388-5398	JJ	denotes	consistent
T10203	5399-5403	IN	denotes	with
T10204	5404-5407	DT	denotes	the
T10205	5408-5415	NN	denotes	ability
T10206	5416-5418	IN	denotes	of
T10207	5419-5423	NN	denotes	BMP2
T10208	5424-5426	TO	denotes	to
T10210	5427-5431	CC	denotes	both
T10209	5432-5440	VB	denotes	increase
T10211	5441-5444	NN	denotes	PKA
T10212	5445-5453	NN	denotes	activity
T10213	5454-5457	CC	denotes	and
T10214	5458-5464	VB	denotes	induce
T10215	5465-5475	NN	denotes	expression
T10216	5476-5478	IN	denotes	of
T10217	5479-5483	NN	denotes	Sox6
T10218	5484-5486	IN	denotes	in
T10219	5487-5493	NN	denotes	tissue
T10221	5494-5501	NN	denotes	culture
T10220	5502-5507	NNS	denotes	cells
T10222	5508-5509	-LRB-	denotes	(
T10223	5509-5512	NNP	denotes	Lee
T10224	5513-5516	CC	denotes	and
T10225	5517-5523	NNP	denotes	Chuong
T10226	5524-5528	CD	denotes	1997
T10227	5528-5529	:	denotes	;
T10228	5530-5539	NNP	denotes	Fernandez
T10229	5539-5540	HYPH	denotes	-
T10230	5540-5546	NNP	denotes	Lloris
T10231	5547-5549	FW	denotes	et
T10232	5550-5553	FW	denotes	al.
T10233	5554-5558	CD	denotes	2003
T10234	5558-5559	-RRB-	denotes	)
T10235	5559-5560	.	denotes	.
T10236	5560-5887	sentence	denotes	Although we have tried to monitor the expression of L-Sox5 or Sox6 in postnatal articular cartilage, and test the phosphorylation state of SOX9 using previously described reagents (Lefebvre et al. 1998; Huang et al. 2000), we have been unable to obtain specific signal at the late postnatal stages required (unpublished data).
T10237	5561-5569	IN	denotes	Although
T10239	5570-5572	PRP	denotes	we
T10240	5573-5577	VBP	denotes	have
T10238	5578-5583	VBN	denotes	tried
T10242	5584-5586	TO	denotes	to
T10243	5587-5594	VB	denotes	monitor
T10244	5595-5598	DT	denotes	the
T10245	5599-5609	NN	denotes	expression
T10246	5610-5612	IN	denotes	of
T10247	5613-5614	NN	denotes	L
T10249	5614-5615	HYPH	denotes	-
T10248	5615-5619	NN	denotes	Sox5
T10250	5620-5622	CC	denotes	or
T10251	5623-5627	NN	denotes	Sox6
T10252	5628-5630	IN	denotes	in
T10253	5631-5640	JJ	denotes	postnatal
T10255	5641-5650	JJ	denotes	articular
T10254	5651-5660	NN	denotes	cartilage
T10256	5660-5662	,	denotes	,
T10257	5662-5665	CC	denotes	and
T10258	5666-5670	VB	denotes	test
T10259	5671-5674	DT	denotes	the
T10261	5675-5690	NN	denotes	phosphorylation
T10260	5691-5696	NN	denotes	state
T10262	5697-5699	IN	denotes	of
T10263	5700-5704	NN	denotes	SOX9
T10264	5705-5710	VBG	denotes	using
T10265	5711-5721	RB	denotes	previously
T10266	5722-5731	VBN	denotes	described
T10267	5732-5740	NNS	denotes	reagents
T10268	5741-5742	-LRB-	denotes	(
T10269	5742-5750	NNP	denotes	Lefebvre
T10270	5751-5753	FW	denotes	et
T10271	5754-5757	FW	denotes	al.
T10272	5758-5762	CD	denotes	1998
T10273	5762-5763	:	denotes	;
T10274	5764-5769	NNP	denotes	Huang
T10275	5770-5772	FW	denotes	et
T10276	5773-5776	FW	denotes	al.
T10277	5777-5781	CD	denotes	2000
T10278	5781-5782	-RRB-	denotes	)
T10279	5782-5784	,	denotes	,
T10280	5784-5786	PRP	denotes	we
T10281	5787-5791	VBP	denotes	have
T10241	5792-5796	VBN	denotes	been
T10282	5797-5803	JJ	denotes	unable
T10283	5804-5806	TO	denotes	to
T10284	5807-5813	VB	denotes	obtain
T10285	5814-5822	JJ	denotes	specific
T10286	5823-5829	NN	denotes	signal
T10287	5830-5832	IN	denotes	at
T10288	5833-5836	DT	denotes	the
T10290	5837-5841	JJ	denotes	late
T10291	5842-5851	JJ	denotes	postnatal
T10289	5852-5858	NNS	denotes	stages
T10292	5859-5867	VBN	denotes	required
T10293	5868-5869	-LRB-	denotes	(
T10295	5869-5880	JJ	denotes	unpublished
T10294	5881-5885	NNS	denotes	data
T10296	5885-5886	-RRB-	denotes	)
T10297	5886-5887	.	denotes	.
T10298	5887-6052	sentence	denotes	Furthermore, null mutations in L-Sox5 or Sox-6 cause lethality at or soon after birth, and no effect on cartilage maintenance has been reported (Smits et al. 2001).
T10299	5888-5899	RB	denotes	Furthermore
T10301	5899-5901	,	denotes	,
T10302	5901-5905	JJ	denotes	null
T10303	5906-5915	NNS	denotes	mutations
T10304	5916-5918	IN	denotes	in
T10305	5919-5920	NN	denotes	L
T10307	5920-5921	HYPH	denotes	-
T10306	5921-5925	NN	denotes	Sox5
T10308	5926-5928	CC	denotes	or
T10309	5929-5932	NN	denotes	Sox
T10310	5932-5933	HYPH	denotes	-
T10311	5933-5934	CD	denotes	6
T10300	5935-5940	VBP	denotes	cause
T10312	5941-5950	NN	denotes	lethality
T10313	5951-5953	IN	denotes	at
T10314	5954-5956	CC	denotes	or
T10315	5957-5961	RB	denotes	soon
T10316	5962-5967	IN	denotes	after
T10317	5968-5973	NN	denotes	birth
T10318	5973-5975	,	denotes	,
T10319	5975-5978	CC	denotes	and
T10320	5979-5981	DT	denotes	no
T10321	5982-5988	NN	denotes	effect
T10323	5989-5991	IN	denotes	on
T10324	5992-6001	NN	denotes	cartilage
T10325	6002-6013	NN	denotes	maintenance
T10326	6014-6017	VBZ	denotes	has
T10327	6018-6022	VBN	denotes	been
T10322	6023-6031	VBN	denotes	reported
T10328	6032-6033	-LRB-	denotes	(
T10329	6033-6038	NNP	denotes	Smits
T10330	6039-6041	FW	denotes	et
T10331	6042-6045	FW	denotes	al.
T10332	6046-6050	CD	denotes	2001
T10333	6050-6051	-RRB-	denotes	)
T10334	6051-6052	.	denotes	.
T10335	6052-6201	sentence	denotes	However, it seems likely that these or other processes regulated by BMP signaling cooperate with SOX9 to induce target genes in articular cartilage.
T10336	6053-6060	RB	denotes	However
T10338	6060-6062	,	denotes	,
T10339	6062-6064	PRP	denotes	it
T10337	6065-6070	VBZ	denotes	seems
T10340	6071-6077	JJ	denotes	likely
T10341	6078-6082	IN	denotes	that
T10343	6083-6088	DT	denotes	these
T10344	6089-6091	CC	denotes	or
T10345	6092-6097	JJ	denotes	other
T10346	6098-6107	NNS	denotes	processes
T10347	6108-6117	VBN	denotes	regulated
T10348	6118-6120	IN	denotes	by
T10349	6121-6124	NN	denotes	BMP
T10350	6125-6134	NN	denotes	signaling
T10342	6135-6144	VBP	denotes	cooperate
T10351	6145-6149	IN	denotes	with
T10352	6150-6154	NN	denotes	SOX9
T10353	6155-6157	TO	denotes	to
T10354	6158-6164	VB	denotes	induce
T10355	6165-6171	JJ	denotes	target
T10356	6172-6177	NNS	denotes	genes
T10357	6178-6180	IN	denotes	in
T10358	6181-6190	JJ	denotes	articular
T10359	6191-6200	NN	denotes	cartilage
T10360	6200-6201	.	denotes	.
T10361	6201-6400	sentence	denotes	Mutation of Smad3 or expression of dominant negative transforming growth factor β (TGF-β) type II receptor also disrupts normal articular cartilage maintenance (Serra et al. 1997; Yang et al. 2001).
T10362	6202-6210	NN	denotes	Mutation
T10364	6211-6213	IN	denotes	of
T10365	6214-6219	NN	denotes	Smad3
T10366	6220-6222	CC	denotes	or
T10367	6223-6233	NN	denotes	expression
T10368	6234-6236	IN	denotes	of
T10369	6237-6245	JJ	denotes	dominant
T10371	6246-6254	JJ	denotes	negative
T10372	6255-6267	VBG	denotes	transforming
T10374	6268-6274	NN	denotes	growth
T10375	6275-6281	NN	denotes	factor
T10373	6282-6283	NN	denotes	β
T10376	6284-6285	-LRB-	denotes	(
T10377	6285-6288	NN	denotes	TGF
T10379	6288-6289	HYPH	denotes	-
T10378	6289-6290	NN	denotes	β
T10380	6290-6291	-RRB-	denotes	)
T10381	6292-6296	NN	denotes	type
T10382	6297-6299	CD	denotes	II
T10370	6300-6308	NN	denotes	receptor
T10383	6309-6313	RB	denotes	also
T10363	6314-6322	VBZ	denotes	disrupts
T10384	6323-6329	JJ	denotes	normal
T10386	6330-6339	JJ	denotes	articular
T10387	6340-6349	NN	denotes	cartilage
T10385	6350-6361	NN	denotes	maintenance
T10388	6362-6363	-LRB-	denotes	(
T10389	6363-6368	NNP	denotes	Serra
T10390	6369-6371	FW	denotes	et
T10391	6372-6375	FW	denotes	al.
T10392	6376-6380	CD	denotes	1997
T10393	6380-6381	:	denotes	;
T10394	6382-6386	NNP	denotes	Yang
T10395	6387-6389	FW	denotes	et
T10396	6390-6393	FW	denotes	al.
T10397	6394-6398	CD	denotes	2001
T10398	6398-6399	-RRB-	denotes	)
T10399	6399-6400	.	denotes	.
T10400	6400-6555	sentence	denotes	Both manipulations should disrupt TGFβ rather than BMP signaling, and both manipulations cause articular cartilage to hypertrophy and be replaced by bone.
T10401	6401-6405	DT	denotes	Both
T10402	6406-6419	NNS	denotes	manipulations
T10404	6420-6426	MD	denotes	should
T10403	6427-6434	VB	denotes	disrupt
T10405	6435-6439	NN	denotes	TGFβ
T10406	6440-6446	JJ	denotes	rather
T10407	6447-6451	IN	denotes	than
T10408	6452-6455	NN	denotes	BMP
T10409	6456-6465	NN	denotes	signaling
T10410	6465-6467	,	denotes	,
T10411	6467-6470	CC	denotes	and
T10412	6471-6475	DT	denotes	both
T10413	6476-6489	NNS	denotes	manipulations
T10414	6490-6495	VBP	denotes	cause
T10415	6496-6505	JJ	denotes	articular
T10416	6506-6515	NN	denotes	cartilage
T10418	6516-6518	TO	denotes	to
T10417	6519-6530	VB	denotes	hypertrophy
T10419	6531-6534	CC	denotes	and
T10420	6535-6537	VB	denotes	be
T10421	6538-6546	VBN	denotes	replaced
T10422	6547-6549	IN	denotes	by
T10423	6550-6554	NN	denotes	bone
T10424	6554-6555	.	denotes	.
T10425	6555-6700	sentence	denotes	In contrast, our analysis of Bmpr1a mutant articular cartilage showed a loss of ECM components, but no signs of hypertrophy or bone replacement.
T10426	6556-6558	IN	denotes	In
T10428	6559-6567	NN	denotes	contrast
T10429	6567-6569	,	denotes	,
T10430	6569-6572	PRP$	denotes	our
T10431	6573-6581	NN	denotes	analysis
T10432	6582-6584	IN	denotes	of
T10433	6585-6591	NN	denotes	Bmpr1a
T10434	6592-6598	NN	denotes	mutant
T10436	6599-6608	JJ	denotes	articular
T10435	6609-6618	NN	denotes	cartilage
T10427	6619-6625	VBD	denotes	showed
T10437	6626-6627	DT	denotes	a
T10438	6628-6632	NN	denotes	loss
T10439	6633-6635	IN	denotes	of
T10440	6636-6639	NN	denotes	ECM
T10441	6640-6650	NNS	denotes	components
T10442	6650-6652	,	denotes	,
T10443	6652-6655	CC	denotes	but
T10444	6656-6658	DT	denotes	no
T10445	6659-6664	NNS	denotes	signs
T10446	6665-6667	IN	denotes	of
T10447	6668-6679	NN	denotes	hypertrophy
T10448	6680-6682	CC	denotes	or
T10449	6683-6687	NN	denotes	bone
T10450	6688-6699	NN	denotes	replacement
T10451	6699-6700	.	denotes	.
T10452	6700-6808	sentence	denotes	Therefore, TGFβ and BMP signaling are playing distinct but necessary roles to maintain articular cartilage.
T10453	6701-6710	RB	denotes	Therefore
T10455	6710-6712	,	denotes	,
T10456	6712-6716	NN	denotes	TGFβ
T10458	6717-6720	CC	denotes	and
T10459	6721-6724	NN	denotes	BMP
T10457	6725-6734	NN	denotes	signaling
T10460	6735-6738	VBP	denotes	are
T10454	6739-6746	VBG	denotes	playing
T10461	6747-6755	JJ	denotes	distinct
T10463	6756-6759	CC	denotes	but
T10464	6760-6769	JJ	denotes	necessary
T10462	6770-6775	NNS	denotes	roles
T10465	6776-6778	TO	denotes	to
T10466	6779-6787	VB	denotes	maintain
T10467	6788-6797	JJ	denotes	articular
T10468	6798-6807	NN	denotes	cartilage
T10469	6807-6808	.	denotes	.
T10470	6808-7192	sentence	denotes	Although BMPs were originally isolated on the basis of their ability to induce ectopic bone formation, their presence in articular cartilage and strong effect on cartilage formation has stimulated interest in using them to repair or regenerate cartilage defects in adult animals (Chang et al. 1994; Erlacher et al. 1998; Edwards and Francis-West 2001; Chubinskaya and Kuettner 2003).
T10471	6809-6817	IN	denotes	Although
T10473	6818-6822	NNS	denotes	BMPs
T10474	6823-6827	VBD	denotes	were
T10475	6828-6838	RB	denotes	originally
T10472	6839-6847	VBN	denotes	isolated
T10477	6848-6850	IN	denotes	on
T10478	6851-6854	DT	denotes	the
T10479	6855-6860	NN	denotes	basis
T10480	6861-6863	IN	denotes	of
T10481	6864-6869	PRP$	denotes	their
T10482	6870-6877	NN	denotes	ability
T10483	6878-6880	TO	denotes	to
T10484	6881-6887	VB	denotes	induce
T10485	6888-6895	JJ	denotes	ectopic
T10487	6896-6900	NN	denotes	bone
T10486	6901-6910	NN	denotes	formation
T10488	6910-6912	,	denotes	,
T10489	6912-6917	PRP$	denotes	their
T10490	6918-6926	NN	denotes	presence
T10491	6927-6929	IN	denotes	in
T10492	6930-6939	JJ	denotes	articular
T10493	6940-6949	NN	denotes	cartilage
T10494	6950-6953	CC	denotes	and
T10495	6954-6960	JJ	denotes	strong
T10496	6961-6967	NN	denotes	effect
T10497	6968-6970	IN	denotes	on
T10498	6971-6980	NN	denotes	cartilage
T10499	6981-6990	NN	denotes	formation
T10500	6991-6994	VBZ	denotes	has
T10476	6995-7005	VBN	denotes	stimulated
T10501	7006-7014	NN	denotes	interest
T10502	7015-7017	IN	denotes	in
T10503	7018-7023	VBG	denotes	using
T10504	7024-7028	PRP	denotes	them
T10505	7029-7031	TO	denotes	to
T10506	7032-7038	VB	denotes	repair
T10507	7039-7041	CC	denotes	or
T10508	7042-7052	VB	denotes	regenerate
T10509	7053-7062	NN	denotes	cartilage
T10510	7063-7070	NNS	denotes	defects
T10511	7071-7073	IN	denotes	in
T10512	7074-7079	NN	denotes	adult
T10513	7080-7087	NNS	denotes	animals
T10514	7088-7089	-LRB-	denotes	(
T10515	7089-7094	NNP	denotes	Chang
T10516	7095-7097	FW	denotes	et
T10517	7098-7101	FW	denotes	al.
T10518	7102-7106	CD	denotes	1994
T10519	7106-7107	:	denotes	;
T10520	7108-7116	NNP	denotes	Erlacher
T10521	7117-7119	FW	denotes	et
T10522	7120-7123	FW	denotes	al.
T10523	7124-7128	CD	denotes	1998
T10524	7128-7129	:	denotes	;
T10525	7130-7137	NNP	denotes	Edwards
T10526	7138-7141	CC	denotes	and
T10527	7142-7149	NNP	denotes	Francis
T10528	7149-7150	HYPH	denotes	-
T10529	7150-7154	NNP	denotes	West
T10530	7155-7159	CD	denotes	2001
T10531	7159-7160	:	denotes	;
T10532	7161-7172	NNP	denotes	Chubinskaya
T10533	7173-7176	CC	denotes	and
T10534	7177-7185	NNP	denotes	Kuettner
T10535	7186-7190	CD	denotes	2003
T10536	7190-7191	-RRB-	denotes	)
T10537	7191-7192	.	denotes	.
T10538	7192-7506	sentence	denotes	The failure to maintain articular cartilage in the absence of normal BMPR1A function suggests that ligands or small molecule agonists that interact specifically with this receptor subtype may be particularly good candidates for designing new approaches to maintain or heal articular cartilage at postnatal stages.
T10539	7193-7196	DT	denotes	The
T10540	7197-7204	NN	denotes	failure
T10542	7205-7207	TO	denotes	to
T10543	7208-7216	VB	denotes	maintain
T10544	7217-7226	JJ	denotes	articular
T10545	7227-7236	NN	denotes	cartilage
T10546	7237-7239	IN	denotes	in
T10547	7240-7243	DT	denotes	the
T10548	7244-7251	NN	denotes	absence
T10549	7252-7254	IN	denotes	of
T10550	7255-7261	JJ	denotes	normal
T10552	7262-7268	NN	denotes	BMPR1A
T10551	7269-7277	NN	denotes	function
T10541	7278-7286	VBZ	denotes	suggests
T10553	7287-7291	IN	denotes	that
T10555	7292-7299	NNS	denotes	ligands
T10556	7300-7302	CC	denotes	or
T10557	7303-7308	JJ	denotes	small
T10559	7309-7317	NN	denotes	molecule
T10558	7318-7326	NNS	denotes	agonists
T10560	7327-7331	WDT	denotes	that
T10561	7332-7340	VBP	denotes	interact
T10562	7341-7353	RB	denotes	specifically
T10563	7354-7358	IN	denotes	with
T10564	7359-7363	DT	denotes	this
T10566	7364-7372	NN	denotes	receptor
T10565	7373-7380	NN	denotes	subtype
T10567	7381-7384	MD	denotes	may
T10554	7385-7387	VB	denotes	be
T10568	7388-7400	RB	denotes	particularly
T10569	7401-7405	JJ	denotes	good
T10570	7406-7416	NNS	denotes	candidates
T10571	7417-7420	IN	denotes	for
T10572	7421-7430	VBG	denotes	designing
T10573	7431-7434	JJ	denotes	new
T10574	7435-7445	NNS	denotes	approaches
T10575	7446-7448	TO	denotes	to
T10576	7449-7457	VB	denotes	maintain
T10577	7458-7460	CC	denotes	or
T10578	7461-7465	VB	denotes	heal
T10579	7466-7475	JJ	denotes	articular
T10580	7476-7485	NN	denotes	cartilage
T10581	7486-7488	IN	denotes	at
T10582	7489-7498	JJ	denotes	postnatal
T10583	7499-7505	NNS	denotes	stages
T10584	7505-7506	.	denotes	.
T10585	7506-7637	sentence	denotes	Lack of Bmpr1a function in articular cartilage results in severe fibrillation of the articular surface and loss of joint mobility.
T10586	7507-7511	NN	denotes	Lack
T10588	7512-7514	IN	denotes	of
T10589	7515-7521	NN	denotes	Bmpr1a
T10590	7522-7530	NN	denotes	function
T10591	7531-7533	IN	denotes	in
T10592	7534-7543	JJ	denotes	articular
T10593	7544-7553	NN	denotes	cartilage
T10587	7554-7561	VBZ	denotes	results
T10594	7562-7564	IN	denotes	in
T10595	7565-7571	JJ	denotes	severe
T10596	7572-7584	NN	denotes	fibrillation
T10597	7585-7587	IN	denotes	of
T10598	7588-7591	DT	denotes	the
T10600	7592-7601	JJ	denotes	articular
T10599	7602-7609	NN	denotes	surface
T10601	7610-7613	CC	denotes	and
T10602	7614-7618	NN	denotes	loss
T10603	7619-7621	IN	denotes	of
T10604	7622-7627	NN	denotes	joint
T10605	7628-7636	NN	denotes	mobility
T10606	7636-7637	.	denotes	.
T10607	7637-7800	sentence	denotes	The development of severe arthritis symptoms in Bmpr1a-deficient mice raises the possibility that defects in BMP signaling also contribute to human joint disease.
T10608	7638-7641	DT	denotes	The
T10609	7642-7653	NN	denotes	development
T10611	7654-7656	IN	denotes	of
T10612	7657-7663	JJ	denotes	severe
T10614	7664-7673	NN	denotes	arthritis
T10613	7674-7682	NNS	denotes	symptoms
T10615	7683-7685	IN	denotes	in
T10616	7686-7692	NN	denotes	Bmpr1a
T10618	7692-7693	HYPH	denotes	-
T10617	7693-7702	JJ	denotes	deficient
T10619	7703-7707	NNS	denotes	mice
T10610	7708-7714	VBZ	denotes	raises
T10620	7715-7718	DT	denotes	the
T10621	7719-7730	NN	denotes	possibility
T10622	7731-7735	IN	denotes	that
T10624	7736-7743	NNS	denotes	defects
T10625	7744-7746	IN	denotes	in
T10626	7747-7750	NN	denotes	BMP
T10627	7751-7760	NN	denotes	signaling
T10628	7761-7765	RB	denotes	also
T10623	7766-7776	VBP	denotes	contribute
T10629	7777-7779	IN	denotes	to
T10630	7780-7785	JJ	denotes	human
T10632	7786-7791	NN	denotes	joint
T10631	7792-7799	NN	denotes	disease
T10633	7799-7800	.	denotes	.
T10634	7800-8013	sentence	denotes	Osteoarthritis is known to have a significant genetic component, but it likely involves multiple genetic factors that have been difficult to identify (Spector et al. 1996; Felson et al. 1998; Hirsch et al. 1998).
T10635	7801-7815	NN	denotes	Osteoarthritis
T10637	7816-7818	VBZ	denotes	is
T10636	7819-7824	VBN	denotes	known
T10638	7825-7827	TO	denotes	to
T10639	7828-7832	VB	denotes	have
T10640	7833-7834	DT	denotes	a
T10642	7835-7846	JJ	denotes	significant
T10643	7847-7854	JJ	denotes	genetic
T10641	7855-7864	NN	denotes	component
T10644	7864-7866	,	denotes	,
T10645	7866-7869	CC	denotes	but
T10646	7870-7872	PRP	denotes	it
T10648	7873-7879	RB	denotes	likely
T10647	7880-7888	VBZ	denotes	involves
T10649	7889-7897	JJ	denotes	multiple
T10651	7898-7905	JJ	denotes	genetic
T10650	7906-7913	NNS	denotes	factors
T10652	7914-7918	WDT	denotes	that
T10654	7919-7923	VBP	denotes	have
T10653	7924-7928	VBN	denotes	been
T10655	7929-7938	JJ	denotes	difficult
T10656	7939-7941	TO	denotes	to
T10657	7942-7950	VB	denotes	identify
T10658	7951-7952	-LRB-	denotes	(
T10659	7952-7959	NNP	denotes	Spector
T10660	7960-7962	FW	denotes	et
T10661	7963-7966	FW	denotes	al.
T10662	7967-7971	CD	denotes	1996
T10663	7971-7972	:	denotes	;
T10664	7973-7979	NNP	denotes	Felson
T10665	7980-7982	FW	denotes	et
T10666	7983-7986	FW	denotes	al.
T10667	7987-7991	CD	denotes	1998
T10668	7991-7992	:	denotes	;
T10669	7993-7999	NNP	denotes	Hirsch
T10670	8000-8002	FW	denotes	et
T10671	8003-8006	FW	denotes	al.
T10672	8007-8011	CD	denotes	1998
T10673	8011-8012	-RRB-	denotes	)
T10674	8012-8013	.	denotes	.
T10675	8013-8239	sentence	denotes	Humans that are heterozygous for loss-of-function mutations in BMPR1A are known to be at risk for juvenile polyposis (Howe et al. 2001; Zhou et al. 2001), but the risk of osteoarthritis for these people has not been reported.
T10676	8014-8020	NNS	denotes	Humans
T10678	8021-8025	WDT	denotes	that
T10679	8026-8029	VBP	denotes	are
T10680	8030-8042	JJ	denotes	heterozygous
T10681	8043-8046	IN	denotes	for
T10682	8047-8051	NN	denotes	loss
T10684	8051-8052	HYPH	denotes	-
T10685	8052-8054	IN	denotes	of
T10686	8054-8055	HYPH	denotes	-
T10687	8055-8063	NN	denotes	function
T10683	8064-8073	NNS	denotes	mutations
T10688	8074-8076	IN	denotes	in
T10689	8077-8083	NN	denotes	BMPR1A
T10690	8084-8087	VBP	denotes	are
T10677	8088-8093	VBN	denotes	known
T10691	8094-8096	TO	denotes	to
T10692	8097-8099	VB	denotes	be
T10693	8100-8102	IN	denotes	at
T10694	8103-8107	NN	denotes	risk
T10695	8108-8111	IN	denotes	for
T10696	8112-8120	JJ	denotes	juvenile
T10697	8121-8130	NN	denotes	polyposis
T10698	8131-8132	-LRB-	denotes	(
T10699	8132-8136	NNP	denotes	Howe
T10700	8137-8139	FW	denotes	et
T10701	8140-8143	FW	denotes	al.
T10702	8144-8148	CD	denotes	2001
T10703	8148-8149	:	denotes	;
T10704	8150-8154	NNP	denotes	Zhou
T10705	8155-8157	FW	denotes	et
T10706	8158-8161	FW	denotes	al.
T10707	8162-8166	CD	denotes	2001
T10708	8166-8167	-RRB-	denotes	)
T10709	8167-8169	,	denotes	,
T10710	8169-8172	CC	denotes	but
T10711	8173-8176	DT	denotes	the
T10712	8177-8181	NN	denotes	risk
T10714	8182-8184	IN	denotes	of
T10715	8185-8199	NN	denotes	osteoarthritis
T10716	8200-8203	IN	denotes	for
T10717	8204-8209	DT	denotes	these
T10718	8210-8216	NNS	denotes	people
T10719	8217-8220	VBZ	denotes	has
T10720	8221-8224	RB	denotes	not
T10721	8225-8229	VBN	denotes	been
T10713	8230-8238	VBN	denotes	reported
T10722	8238-8239	.	denotes	.
T10723	8239-8396	sentence	denotes	However, the control mice used in this study were heterozygous for a null allele of Bmpr1a, and they showed little sign of osteoarthritis even late in life.
T10724	8240-8247	RB	denotes	However
T10726	8247-8249	,	denotes	,
T10727	8249-8252	DT	denotes	the
T10729	8253-8260	NN	denotes	control
T10728	8261-8265	NNS	denotes	mice
T10730	8266-8270	VBN	denotes	used
T10731	8271-8273	IN	denotes	in
T10732	8274-8278	DT	denotes	this
T10733	8279-8284	NN	denotes	study
T10725	8285-8289	VBD	denotes	were
T10734	8290-8302	JJ	denotes	heterozygous
T10735	8303-8306	IN	denotes	for
T10736	8307-8308	DT	denotes	a
T10738	8309-8313	JJ	denotes	null
T10737	8314-8320	NN	denotes	allele
T10739	8321-8323	IN	denotes	of
T10740	8324-8330	NN	denotes	Bmpr1a
T10741	8330-8332	,	denotes	,
T10742	8332-8335	CC	denotes	and
T10743	8336-8340	PRP	denotes	they
T10744	8341-8347	VBD	denotes	showed
T10745	8348-8354	JJ	denotes	little
T10746	8355-8359	NN	denotes	sign
T10747	8360-8362	IN	denotes	of
T10748	8363-8377	NN	denotes	osteoarthritis
T10749	8378-8382	RB	denotes	even
T10750	8383-8387	RB	denotes	late
T10751	8388-8390	IN	denotes	in
T10752	8391-8395	NN	denotes	life
T10753	8395-8396	.	denotes	.
T10754	8396-8565	sentence	denotes	Several chromosome regions have been previously linked to arthritis phenotypes in humans using either association studies in populations or linkage studies in families.
T10755	8397-8404	JJ	denotes	Several
T10757	8405-8415	NN	denotes	chromosome
T10756	8416-8423	NNS	denotes	regions
T10759	8424-8428	VBP	denotes	have
T10760	8429-8433	VBN	denotes	been
T10761	8434-8444	RB	denotes	previously
T10758	8445-8451	VBN	denotes	linked
T10762	8452-8454	IN	denotes	to
T10763	8455-8464	NN	denotes	arthritis
T10764	8465-8475	NNS	denotes	phenotypes
T10765	8476-8478	IN	denotes	in
T10766	8479-8485	NNS	denotes	humans
T10767	8486-8491	VBG	denotes	using
T10768	8492-8498	CC	denotes	either
T10770	8499-8510	NN	denotes	association
T10769	8511-8518	NNS	denotes	studies
T10771	8519-8521	IN	denotes	in
T10772	8522-8533	NNS	denotes	populations
T10773	8534-8536	CC	denotes	or
T10774	8537-8544	NN	denotes	linkage
T10775	8545-8552	NNS	denotes	studies
T10776	8553-8555	IN	denotes	in
T10777	8556-8564	NNS	denotes	families
T10778	8564-8565	.	denotes	.
T10779	8565-8935	sentence	denotes	It is interesting to note that several of these chromosome regions contain genes encoding different members of the BMP signaling pathway, including the BMP5 gene on human chromosome 6p12 (Loughlin et al. 2002), the MADH1 gene on human chromosome 4q26–4q31 (Leppavuori et al. 1999; Kent et al. 2002), and the BMPR2 receptor on human chromosome 2q33 (Wright et al. 1996).
T10780	8566-8568	PRP	denotes	It
T10781	8569-8571	VBZ	denotes	is
T10782	8572-8583	JJ	denotes	interesting
T10783	8584-8586	TO	denotes	to
T10784	8587-8591	VB	denotes	note
T10785	8592-8596	IN	denotes	that
T10787	8597-8604	JJ	denotes	several
T10788	8605-8607	IN	denotes	of
T10789	8608-8613	DT	denotes	these
T10791	8614-8624	NN	denotes	chromosome
T10790	8625-8632	NNS	denotes	regions
T10786	8633-8640	VBP	denotes	contain
T10792	8641-8646	NNS	denotes	genes
T10793	8647-8655	VBG	denotes	encoding
T10794	8656-8665	JJ	denotes	different
T10795	8666-8673	NNS	denotes	members
T10796	8674-8676	IN	denotes	of
T10797	8677-8680	DT	denotes	the
T10799	8681-8684	NN	denotes	BMP
T10800	8685-8694	NN	denotes	signaling
T10798	8695-8702	NN	denotes	pathway
T10801	8702-8704	,	denotes	,
T10802	8704-8713	VBG	denotes	including
T10803	8714-8717	DT	denotes	the
T10805	8718-8722	NN	denotes	BMP5
T10804	8723-8727	NN	denotes	gene
T10806	8728-8730	IN	denotes	on
T10807	8731-8736	JJ	denotes	human
T10809	8737-8747	NN	denotes	chromosome
T10808	8748-8752	NN	denotes	6p12
T10810	8753-8754	-LRB-	denotes	(
T10811	8754-8762	NNP	denotes	Loughlin
T10812	8763-8765	FW	denotes	et
T10813	8766-8769	FW	denotes	al.
T10814	8770-8774	CD	denotes	2002
T10815	8774-8775	-RRB-	denotes	)
T10816	8775-8777	,	denotes	,
T10817	8777-8780	DT	denotes	the
T10819	8781-8786	NN	denotes	MADH1
T10818	8787-8791	NN	denotes	gene
T10820	8792-8794	IN	denotes	on
T10821	8795-8800	JJ	denotes	human
T10823	8801-8811	NN	denotes	chromosome
T10822	8812-8816	NN	denotes	4q26
T10824	8816-8817	SYM	denotes	–
T10825	8817-8821	NN	denotes	4q31
T10826	8822-8823	-LRB-	denotes	(
T10827	8823-8833	NNP	denotes	Leppavuori
T10828	8834-8836	FW	denotes	et
T10829	8837-8840	FW	denotes	al.
T10830	8841-8845	CD	denotes	1999
T10831	8845-8846	:	denotes	;
T10832	8847-8851	NNP	denotes	Kent
T10833	8852-8854	FW	denotes	et
T10834	8855-8858	FW	denotes	al.
T10835	8859-8863	CD	denotes	2002
T10836	8863-8864	-RRB-	denotes	)
T10837	8864-8866	,	denotes	,
T10838	8866-8869	CC	denotes	and
T10839	8870-8873	DT	denotes	the
T10841	8874-8879	NN	denotes	BMPR2
T10840	8880-8888	NN	denotes	receptor
T10842	8889-8891	IN	denotes	on
T10843	8892-8897	JJ	denotes	human
T10845	8898-8908	NN	denotes	chromosome
T10844	8909-8913	NN	denotes	2q33
T10846	8914-8915	-LRB-	denotes	(
T10847	8915-8921	NNP	denotes	Wright
T10848	8922-8924	FW	denotes	et
T10849	8925-8928	FW	denotes	al.
T10850	8929-8933	CD	denotes	1996
T10851	8933-8934	-RRB-	denotes	)
T10852	8934-8935	.	denotes	.
T10853	8935-9088	sentence	denotes	The complex nature of human osteoarthritis suggests that interactions between multiple genes may be involved in modifying susceptibility to the disease.
T10854	8936-8939	DT	denotes	The
T10856	8940-8947	JJ	denotes	complex
T10855	8948-8954	NN	denotes	nature
T10858	8955-8957	IN	denotes	of
T10859	8958-8963	JJ	denotes	human
T10860	8964-8978	NN	denotes	osteoarthritis
T10857	8979-8987	VBZ	denotes	suggests
T10861	8988-8992	IN	denotes	that
T10863	8993-9005	NNS	denotes	interactions
T10864	9006-9013	IN	denotes	between
T10865	9014-9022	JJ	denotes	multiple
T10866	9023-9028	NNS	denotes	genes
T10867	9029-9032	MD	denotes	may
T10868	9033-9035	VB	denotes	be
T10862	9036-9044	VBN	denotes	involved
T10869	9045-9047	IN	denotes	in
T10870	9048-9057	VBG	denotes	modifying
T10871	9058-9072	NN	denotes	susceptibility
T10872	9073-9075	IN	denotes	to
T10873	9076-9079	DT	denotes	the
T10874	9080-9087	NN	denotes	disease
T10875	9087-9088	.	denotes	.
T10876	9088-9266	sentence	denotes	The inclusion of genetic markers near BMP signaling components may help identify additional osteoarthritis susceptibility loci and facilitate the search for causative mutations.
T10877	9089-9092	DT	denotes	The
T10878	9093-9102	NN	denotes	inclusion
T10880	9103-9105	IN	denotes	of
T10881	9106-9113	JJ	denotes	genetic
T10882	9114-9121	NNS	denotes	markers
T10883	9122-9126	IN	denotes	near
T10884	9127-9130	NN	denotes	BMP
T10886	9131-9140	NN	denotes	signaling
T10885	9141-9151	NNS	denotes	components
T10887	9152-9155	MD	denotes	may
T10879	9156-9160	VB	denotes	help
T10888	9161-9169	VB	denotes	identify
T10889	9170-9180	JJ	denotes	additional
T10891	9181-9195	NN	denotes	osteoarthritis
T10892	9196-9210	NN	denotes	susceptibility
T10890	9211-9215	NNS	denotes	loci
T10893	9216-9219	CC	denotes	and
T10894	9220-9230	VB	denotes	facilitate
T10895	9231-9234	DT	denotes	the
T10896	9235-9241	NN	denotes	search
T10897	9242-9245	IN	denotes	for
T10898	9246-9255	JJ	denotes	causative
T10899	9256-9265	NNS	denotes	mutations
T10900	9265-9266	.	denotes	.
T10901	9266-9465	sentence	denotes	Development and disease processes in synovial joints have been difficult to study genetically, because synovial joints are generated and function at relatively late stages of vertebrate development.
T10902	9267-9278	NN	denotes	Development
T10904	9279-9282	CC	denotes	and
T10905	9283-9290	NN	denotes	disease
T10903	9291-9300	NNS	denotes	processes
T10907	9301-9303	IN	denotes	in
T10908	9304-9312	JJ	denotes	synovial
T10909	9313-9319	NNS	denotes	joints
T10910	9320-9324	VBP	denotes	have
T10906	9325-9329	VBN	denotes	been
T10911	9330-9339	JJ	denotes	difficult
T10912	9340-9342	TO	denotes	to
T10913	9343-9348	VB	denotes	study
T10914	9349-9360	RB	denotes	genetically
T10915	9360-9362	,	denotes	,
T10916	9362-9369	IN	denotes	because
T10918	9370-9378	JJ	denotes	synovial
T10919	9379-9385	NNS	denotes	joints
T10920	9386-9389	VBP	denotes	are
T10917	9390-9399	VBN	denotes	generated
T10921	9400-9403	CC	denotes	and
T10922	9404-9412	VBP	denotes	function
T10923	9413-9415	IN	denotes	at
T10924	9416-9426	RB	denotes	relatively
T10925	9427-9431	JJ	denotes	late
T10926	9432-9438	NNS	denotes	stages
T10927	9439-9441	IN	denotes	of
T10928	9442-9452	NN	denotes	vertebrate
T10929	9453-9464	NN	denotes	development
T10930	9464-9465	.	denotes	.
T10931	9465-9659	sentence	denotes	The Gdf5-Cre system provides a new method for restricting gene expression or inactivation primarily to articular regions, thus avoiding the pleiotropic functions of many genes in other tissues.
T10932	9466-9469	DT	denotes	The
T10934	9470-9474	NN	denotes	Gdf5
T10936	9474-9475	HYPH	denotes	-
T10935	9475-9478	NN	denotes	Cre
T10933	9479-9485	NN	denotes	system
T10937	9486-9494	VBZ	denotes	provides
T10938	9495-9496	DT	denotes	a
T10940	9497-9500	JJ	denotes	new
T10939	9501-9507	NN	denotes	method
T10941	9508-9511	IN	denotes	for
T10942	9512-9523	VBG	denotes	restricting
T10943	9524-9528	NN	denotes	gene
T10944	9529-9539	NN	denotes	expression
T10945	9540-9542	CC	denotes	or
T10946	9543-9555	NN	denotes	inactivation
T10947	9556-9565	RB	denotes	primarily
T10948	9566-9568	IN	denotes	to
T10949	9569-9578	JJ	denotes	articular
T10950	9579-9586	NNS	denotes	regions
T10951	9586-9588	,	denotes	,
T10952	9588-9592	RB	denotes	thus
T10953	9593-9601	VBG	denotes	avoiding
T10954	9602-9605	DT	denotes	the
T10956	9606-9617	JJ	denotes	pleiotropic
T10955	9618-9627	NNS	denotes	functions
T10957	9628-9630	IN	denotes	of
T10958	9631-9635	JJ	denotes	many
T10959	9636-9641	NNS	denotes	genes
T10960	9642-9644	IN	denotes	in
T10961	9645-9650	JJ	denotes	other
T10962	9651-9658	NNS	denotes	tissues
T10963	9658-9659	.	denotes	.
T10964	9659-9905	sentence	denotes	Depending on the configuration of the floxed target gene, this system can be used to either activate the expression of a gene primarily in developing joints (ssee Figure 1B–1D), or to inactivate gene function in articular regions (see Figure 3).
T10965	9660-9669	VBG	denotes	Depending
T10967	9670-9672	IN	denotes	on
T10968	9673-9676	DT	denotes	the
T10969	9677-9690	NN	denotes	configuration
T10970	9691-9693	IN	denotes	of
T10971	9694-9697	DT	denotes	the
T10973	9698-9704	VBN	denotes	floxed
T10974	9705-9711	NN	denotes	target
T10972	9712-9716	NN	denotes	gene
T10975	9716-9718	,	denotes	,
T10976	9718-9722	DT	denotes	this
T10977	9723-9729	NN	denotes	system
T10978	9730-9733	MD	denotes	can
T10979	9734-9736	VB	denotes	be
T10966	9737-9741	VBN	denotes	used
T10980	9742-9744	TO	denotes	to
T10982	9745-9751	CC	denotes	either
T10981	9752-9760	VB	denotes	activate
T10983	9761-9764	DT	denotes	the
T10984	9765-9775	NN	denotes	expression
T10985	9776-9778	IN	denotes	of
T10986	9779-9780	DT	denotes	a
T10987	9781-9785	NN	denotes	gene
T10988	9786-9795	RB	denotes	primarily
T10989	9796-9798	IN	denotes	in
T10990	9799-9809	VBG	denotes	developing
T10991	9810-9816	NNS	denotes	joints
T10992	9817-9818	-LRB-	denotes	(
T10993	9818-9822	VB	denotes	ssee
T10994	9823-9829	NN	denotes	Figure
T10995	9830-9832	CD	denotes	1B
T10996	9832-9833	SYM	denotes	–
T10997	9833-9835	CD	denotes	1D
T10998	9835-9836	-RRB-	denotes	)
T10999	9836-9838	,	denotes	,
T11000	9838-9840	CC	denotes	or
T11001	9841-9843	TO	denotes	to
T11002	9844-9854	VB	denotes	inactivate
T11003	9855-9859	NN	denotes	gene
T11004	9860-9868	NN	denotes	function
T11005	9869-9871	IN	denotes	in
T11006	9872-9881	JJ	denotes	articular
T11007	9882-9889	NNS	denotes	regions
T11008	9890-9891	-LRB-	denotes	(
T11009	9891-9894	VB	denotes	see
T11010	9895-9901	NN	denotes	Figure
T11011	9902-9903	CD	denotes	3
T11012	9903-9904	-RRB-	denotes	)
T11013	9904-9905	.	denotes	.
T11014	9905-10120	sentence	denotes	Additional studies with this system should greatly enhance our knowledge of the development, function, and disease mechanisms of joints, and may bring us closer to better prevention and treatment of joint diseases.
T11015	9906-9916	JJ	denotes	Additional
T11016	9917-9924	NNS	denotes	studies
T11018	9925-9929	IN	denotes	with
T11019	9930-9934	DT	denotes	this
T11020	9935-9941	NN	denotes	system
T11021	9942-9948	MD	denotes	should
T11022	9949-9956	RB	denotes	greatly
T11017	9957-9964	VB	denotes	enhance
T11023	9965-9968	PRP$	denotes	our
T11024	9969-9978	NN	denotes	knowledge
T11025	9979-9981	IN	denotes	of
T11026	9982-9985	DT	denotes	the
T11028	9986-9997	NN	denotes	development
T11029	9997-9999	,	denotes	,
T11030	9999-10007	NN	denotes	function
T11031	10007-10009	,	denotes	,
T11032	10009-10012	CC	denotes	and
T11033	10013-10020	NN	denotes	disease
T11027	10021-10031	NNS	denotes	mechanisms
T11034	10032-10034	IN	denotes	of
T11035	10035-10041	NNS	denotes	joints
T11036	10041-10043	,	denotes	,
T11037	10043-10046	CC	denotes	and
T11038	10047-10050	MD	denotes	may
T11039	10051-10056	VB	denotes	bring
T11040	10057-10059	PRP	denotes	us
T11041	10060-10066	JJR	denotes	closer
T11042	10067-10069	IN	denotes	to
T11043	10070-10076	JJR	denotes	better
T11044	10077-10087	NN	denotes	prevention
T11045	10088-10091	CC	denotes	and
T11046	10092-10101	NN	denotes	treatment
T11047	10102-10104	IN	denotes	of
T11048	10105-10110	NN	denotes	joint
T11049	10111-10119	NNS	denotes	diseases
T11050	10119-10120	.	denotes	.
R5531	T9223	T9224	amod	Previous,studies
R5532	T9224	T9225	nsubj	studies,suggest
R5533	T9226	T9227	mark	that,involved
R5534	T9227	T9225	ccomp	involved,suggest
R5535	T9228	T9229	compound	BMP,signaling
R5536	T9229	T9227	nsubjpass	signaling,involved
R5537	T9230	T9227	auxpass	is,involved
R5538	T9231	T9227	prep	in,involved
R5539	T9232	T9233	det	a,number
R5540	T9233	T9231	pobj	number,in
R5541	T9234	T9233	amod	large,number
R5542	T9235	T9233	prep	of,number
R5543	T9236	T9237	amod	developmental,events
R5544	T9237	T9235	pobj	events,of
R5545	T9238	T9225	punct	.,suggest
R5546	T9240	T9241	nsubj	Many,occur
R5547	T9242	T9240	prep	of,Many
R5548	T9243	T9244	det	these,events
R5549	T9244	T9242	pobj	events,of
R5550	T9245	T9241	advmod	early,occur
R5551	T9246	T9241	prep	in,occur
R5552	T9247	T9246	pobj	embryogenesis,in
R5553	T9248	T9241	punct	", ",occur
R5554	T9249	T9241	cc	and,occur
R5555	T9250	T9251	amod	complete,inactivation
R5556	T9251	T9252	nsubj	inactivation,causes
R5557	T9252	T9241	conj	causes,occur
R5558	T9253	T9251	prep	of,inactivation
R5559	T9254	T9255	compound	BMP,receptors
R5560	T9255	T9253	pobj	receptors,of
R5561	T9256	T9252	dobj	death,causes
R5562	T9257	T9252	prep	by,causes
R5563	T9258	T9257	pobj	E9.5,by
R5564	T9259	T9260	punct	(,Mishina
R5565	T9260	T9252	meta	Mishina,causes
R5566	T9261	T9260	nmod	et,Mishina
R5567	T9262	T9260	nmod	al.,Mishina
R5568	T9263	T9260	nummod	1995,Mishina
R5569	T9264	T9260	punct	),Mishina
R5570	T9265	T9252	punct	.,causes
R5571	T9267	T9268	det	The,system
R5572	T9268	T9273	nsubj	system,bypasses
R5573	T9269	T9270	compound	Gdf5,Cre
R5574	T9270	T9268	compound	Cre,system
R5575	T9271	T9270	punct	-,Cre
R5576	T9272	T9268	compound	recombination,system
R5577	T9274	T9275	det	the,lethality
R5578	T9275	T9273	dobj	lethality,bypasses
R5579	T9276	T9275	amod	early,lethality
R5580	T9277	T9275	amod	embryonic,lethality
R5581	T9278	T9275	prep	of,lethality
R5582	T9279	T9280	compound	Bmpr1a,mutations
R5583	T9280	T9278	pobj	mutations,of
R5584	T9281	T9273	punct	", ",bypasses
R5585	T9282	T9273	cc	and,bypasses
R5586	T9283	T9273	conj	provides,bypasses
R5587	T9284	T9285	amod	important,information
R5588	T9285	T9283	dobj	information,provides
R5589	T9286	T9285	amod	new,information
R5590	T9287	T9285	prep	about,information
R5591	T9288	T9289	det	the,role
R5592	T9289	T9287	pobj	role,about
R5593	T9290	T9289	prep	of,role
R5594	T9291	T9292	det	this,receptor
R5595	T9292	T9290	pobj	receptor,of
R5596	T9293	T9289	prep	in,role
R5597	T9294	T9295	nmod	limb,development
R5598	T9295	T9293	pobj	development,in
R5599	T9296	T9294	cc	and,limb
R5600	T9297	T9294	conj	skeletal,limb
R5601	T9298	T9273	punct	.,bypasses
R5602	T9300	T9301	det	The,phenotypes
R5603	T9301	T9305	nsubj	phenotypes,include
R5604	T9302	T9301	nummod	three,phenotypes
R5605	T9303	T9301	amod	major,phenotypes
R5606	T9304	T9301	compound	limb,phenotypes
R5607	T9306	T9301	acl	revealed,phenotypes
R5608	T9307	T9306	agent	by,revealed
R5609	T9308	T9307	pcomp	eliminating,by
R5610	T9309	T9308	dobj	Bmpr1a,eliminating
R5611	T9310	T9308	prep	with,eliminating
R5612	T9311	T9312	npadvmod	Gdf5,driven
R5613	T9312	T9314	amod	driven,Cre
R5614	T9313	T9312	punct	-,driven
R5615	T9314	T9310	pobj	Cre,with
R5616	T9315	T9305	dobj	webbing,include
R5617	T9316	T9315	prep	between,webbing
R5618	T9317	T9316	pobj	digits,between
R5619	T9318	T9315	punct	", ",webbing
R5620	T9319	T9315	conj	lack,webbing
R5621	T9320	T9319	prep	of,lack
R5622	T9321	T9322	compound	joint,formation
R5623	T9322	T9320	pobj	formation,of
R5624	T9323	T9319	prep	at,lack
R5625	T9324	T9325	amod	specific,locations
R5626	T9325	T9323	pobj	locations,at
R5627	T9326	T9325	prep	in,locations
R5628	T9327	T9328	det	the,ankle
R5629	T9328	T9326	pobj	ankle,in
R5630	T9329	T9319	punct	", ",lack
R5631	T9330	T9319	cc	and,lack
R5632	T9331	T9319	conj	failure,lack
R5633	T9332	T9333	aux	to,maintain
R5634	T9333	T9331	acl	maintain,failure
R5635	T9334	T9335	amod	articular,cartilage
R5636	T9335	T9333	dobj	cartilage,maintain
R5637	T9336	T9333	prep	after,maintain
R5638	T9337	T9336	pobj	birth,after
R5639	T9338	T9333	punct	", ",maintain
R5640	T9339	T9333	advcl	resulting,maintain
R5641	T9340	T9339	prep	in,resulting
R5642	T9341	T9342	amod	severe,arthritis
R5643	T9342	T9340	pobj	arthritis,in
R5644	T9343	T9305	punct	.,include
R5645	T9345	T9346	amod	Previous,studies
R5646	T9346	T9347	nsubj	studies,shown
R5647	T9348	T9347	aux	have,shown
R5648	T9349	T9350	mark	that,alters
R5649	T9350	T9347	ccomp	alters,shown
R5650	T9351	T9350	nsubj	manipulation,alters
R5651	T9352	T9351	prep	of,manipulation
R5652	T9353	T9354	compound	BMP,signaling
R5653	T9354	T9352	pobj	signaling,of
R5654	T9355	T9356	amod	interdigital,apoptosis
R5655	T9356	T9350	dobj	apoptosis,alters
R5656	T9357	T9350	prep	during,alters
R5657	T9358	T9357	pobj	development,during
R5658	T9359	T9358	prep	of,development
R5659	T9360	T9361	det	the,limb
R5660	T9361	T9359	pobj	limb,of
R5661	T9362	T9347	punct	", ",shown
R5662	T9363	T9347	cc	but,shown
R5663	T9364	T9365	det	no,experiment
R5664	T9365	T9366	nsubj	experiment,identified
R5665	T9366	T9347	conj	identified,shown
R5666	T9367	T9366	aux	has,identified
R5667	T9368	T9369	det	a,member
R5668	T9369	T9366	dobj	member,identified
R5669	T9370	T9369	amod	specific,member
R5670	T9371	T9369	prep	of,member
R5671	T9372	T9373	det	the,pathway
R5672	T9373	T9371	pobj	pathway,of
R5673	T9374	T9375	compound	BMP,signaling
R5674	T9375	T9373	compound	signaling,pathway
R5675	T9376	T9377	dep	that,required
R5676	T9377	T9369	relcl	required,member
R5677	T9378	T9377	auxpass	is,required
R5678	T9379	T9377	prep	for,required
R5679	T9380	T9381	det	this,process
R5680	T9381	T9379	pobj	process,for
R5681	T9382	T9383	punct	(,Yokouchi
R5682	T9383	T9366	meta	Yokouchi,identified
R5683	T9384	T9383	nmod	et,Yokouchi
R5684	T9385	T9383	nmod	al.,Yokouchi
R5685	T9386	T9383	nummod	1996,Yokouchi
R5686	T9387	T9383	punct	;,Yokouchi
R5687	T9388	T9383	conj	Zou,Yokouchi
R5688	T9389	T9388	cc	and,Zou
R5689	T9390	T9388	conj	Niswander,Zou
R5690	T9391	T9390	nummod	1996,Niswander
R5691	T9392	T9390	punct	;,Niswander
R5692	T9393	T9390	conj	Zou,Niswander
R5693	T9394	T9393	nmod	et,Zou
R5694	T9395	T9393	nmod	al.,Zou
R5695	T9396	T9393	nummod	1997,Zou
R5696	T9397	T9393	punct	;,Zou
R5697	T9398	T9393	conj	Guha,Zou
R5698	T9399	T9398	nmod	et,Guha
R5699	T9400	T9398	nmod	al.,Guha
R5700	T9401	T9398	nummod	2002,Guha
R5701	T9402	T9398	punct	),Guha
R5702	T9403	T9366	punct	.,identified
R5703	T9405	T9406	poss	Our,data
R5704	T9406	T9413	nsubj	data,confirm
R5705	T9407	T9406	amod	new,data
R5706	T9408	T9406	nmod	loss,data
R5707	T9409	T9408	punct	-,loss
R5708	T9410	T9408	prep	of,loss
R5709	T9411	T9410	punct	-,of
R5710	T9412	T9410	pobj	function,of
R5711	T9414	T9415	mark	that,required
R5712	T9415	T9413	ccomp	required,confirm
R5713	T9416	T9417	compound	BMP,signaling
R5714	T9417	T9415	nsubjpass	signaling,required
R5715	T9418	T9415	auxpass	is,required
R5716	T9419	T9415	prep	for,required
R5717	T9420	T9421	amod	interdigital,apoptosis
R5718	T9421	T9419	pobj	apoptosis,for
R5719	T9422	T9415	cc	and,required
R5720	T9423	T9415	conj	suggests,required
R5721	T9424	T9425	mark	that,is
R5722	T9425	T9423	ccomp	is,suggests
R5723	T9426	T9425	nsubj	Bmpr1a,is
R5724	T9427	T9428	det	a,component
R5725	T9428	T9425	attr	component,is
R5726	T9429	T9428	amod	critical,component
R5727	T9430	T9428	prep	for,component
R5728	T9431	T9430	pcomp	mediating,for
R5729	T9432	T9433	det	this,signal
R5730	T9433	T9431	dobj	signal,mediating
R5731	T9434	T9413	punct	.,confirm
R5732	T9436	T9437	prep	At,leads
R5733	T9438	T9439	det	some,sites
R5734	T9439	T9436	pobj	sites,At
R5735	T9440	T9437	punct	", ",leads
R5736	T9441	T9437	nsubj	loss,leads
R5737	T9442	T9441	prep	of,loss
R5738	T9443	T9444	compound	Bmpr1a,function
R5739	T9444	T9442	pobj	function,of
R5740	T9445	T9437	prep	to,leads
R5741	T9446	T9447	det	a,defect
R5742	T9447	T9445	pobj	defect,to
R5743	T9448	T9447	prep	in,defect
R5744	T9449	T9450	det	the,stages
R5745	T9450	T9448	pobj	stages,in
R5746	T9451	T9450	amod	early,stages
R5747	T9452	T9450	prep	of,stages
R5748	T9453	T9454	compound	joint,formation
R5749	T9454	T9452	pobj	formation,of
R5750	T9455	T9437	punct	", ",leads
R5751	T9456	T9437	advcl	resulting,leads
R5752	T9457	T9456	prep	in,resulting
R5753	T9458	T9459	det	a,failure
R5754	T9459	T9457	pobj	failure,in
R5755	T9460	T9459	amod	complete,failure
R5756	T9461	T9462	aux	to,form
R5757	T9462	T9459	acl	form,failure
R5758	T9463	T9464	det	a,joint
R5759	T9464	T9462	dobj	joint,form
R5760	T9465	T9459	cc	and,failure
R5761	T9466	T9459	conj	fusion,failure
R5762	T9467	T9466	prep	of,fusion
R5763	T9468	T9467	pobj	bones,of
R5764	T9469	T9466	prep	in,fusion
R5765	T9470	T9471	det	the,ankle
R5766	T9471	T9469	pobj	ankle,in
R5767	T9472	T9437	punct	.,leads
R5768	T9474	T9475	nsubj	Mutations,produce
R5769	T9476	T9474	prep	in,Mutations
R5770	T9477	T9478	nummod	two,ligands
R5771	T9478	T9476	pobj	ligands,in
R5772	T9479	T9478	amod	different,ligands
R5773	T9480	T9478	prep	in,ligands
R5774	T9481	T9482	det	the,family
R5775	T9482	T9480	pobj	family,in
R5776	T9483	T9482	compound	BMP,family
R5777	T9484	T9478	punct	", ",ligands
R5778	T9485	T9478	appos	Gdf5,ligands
R5779	T9486	T9485	cc	and,Gdf5
R5780	T9487	T9485	conj	Gdf6,Gdf5
R5781	T9488	T9478	punct	", ",ligands
R5782	T9489	T9490	det	the,receptor
R5783	T9490	T9478	appos	receptor,ligands
R5784	T9491	T9490	compound	Bmpr1b,receptor
R5785	T9492	T9476	punct	", ",in
R5786	T9493	T9476	cc	and,in
R5787	T9494	T9476	conj	in,in
R5788	T9495	T9496	det	the,locus
R5789	T9496	T9494	pobj	locus,in
R5790	T9497	T9496	amod	human,locus
R5791	T9498	T9496	compound	Noggin,locus
R5792	T9499	T9500	punct	(,Storm
R5793	T9500	T9474	meta	Storm,Mutations
R5794	T9501	T9500	cc	and,Storm
R5795	T9502	T9500	conj	Kingsley,Storm
R5796	T9503	T9502	nummod	1996,Kingsley
R5797	T9504	T9502	punct	;,Kingsley
R5798	T9505	T9502	conj	Gong,Kingsley
R5799	T9506	T9505	nmod	et,Gong
R5800	T9507	T9505	nmod	al.,Gong
R5801	T9508	T9505	nummod	1999,Gong
R5802	T9509	T9505	punct	;,Gong
R5803	T9510	T9505	conj	Baur,Gong
R5804	T9511	T9510	nmod	et,Baur
R5805	T9512	T9510	nmod	al.,Baur
R5806	T9513	T9510	nummod	2000,Baur
R5807	T9514	T9510	punct	;,Baur
R5808	T9515	T9510	conj	Yi,Baur
R5809	T9516	T9515	nmod	et,Yi
R5810	T9517	T9515	nmod	al.,Yi
R5811	T9518	T9515	nummod	2000,Yi
R5812	T9519	T9515	punct	;,Yi
R5813	T9520	T9515	conj	Settle,Yi
R5814	T9521	T9520	nmod	et,Settle
R5815	T9522	T9520	nmod	al.,Settle
R5816	T9523	T9520	nummod	2003,Settle
R5817	T9524	T9520	punct	),Settle
R5818	T9525	T9475	advmod	also,produce
R5819	T9526	T9475	dobj	defects,produce
R5820	T9527	T9526	prep	in,defects
R5821	T9528	T9529	compound	joint,formation
R5822	T9529	T9527	pobj	formation,in
R5823	T9530	T9475	prep	at,produce
R5824	T9531	T9532	amod	specific,locations
R5825	T9532	T9530	pobj	locations,at
R5826	T9533	T9532	prep	in,locations
R5827	T9534	T9535	det	the,limbs
R5828	T9535	T9533	pobj	limbs,in
R5829	T9536	T9475	punct	.,produce
R5830	T9538	T9539	det	The,defects
R5831	T9539	T9541	nsubj	defects,provide
R5832	T9540	T9539	compound	joint,defects
R5833	T9542	T9539	acl	associated,defects
R5834	T9543	T9542	prep	with,associated
R5835	T9544	T9545	amod	multiple,components
R5836	T9545	T9543	pobj	components,with
R5837	T9546	T9545	prep	of,components
R5838	T9547	T9548	det	the,pathway
R5839	T9548	T9546	pobj	pathway,of
R5840	T9549	T9548	compound	BMP,pathway
R5841	T9550	T9551	amod	strong,evidence
R5842	T9551	T9541	dobj	evidence,provide
R5843	T9552	T9553	mark	that,required
R5844	T9553	T9551	acl	required,evidence
R5845	T9554	T9555	compound	BMP,signaling
R5846	T9555	T9553	nsubjpass	signaling,required
R5847	T9556	T9553	auxpass	is,required
R5848	T9557	T9553	prep	for,required
R5849	T9558	T9559	amod	early,stages
R5850	T9559	T9557	pobj	stages,for
R5851	T9560	T9559	prep	of,stages
R5852	T9561	T9562	compound	joint,formation
R5853	T9562	T9560	pobj	formation,of
R5854	T9563	T9553	prep	at,required
R5855	T9564	T9565	det	some,locations
R5856	T9565	T9563	pobj	locations,at
R5857	T9566	T9565	amod	anatomical,locations
R5858	T9567	T9541	punct	.,provide
R5859	T9569	T9570	amod	Most,joints
R5860	T9570	T9571	nsubj	joints,form
R5861	T9572	T9571	advmod	still,form
R5862	T9573	T9571	advmod	normally,form
R5863	T9574	T9575	advmod	when,knocked
R5864	T9575	T9571	advcl	knocked,form
R5865	T9576	T9575	nsubjpass	Bmpr1a,knocked
R5866	T9577	T9575	auxpass	is,knocked
R5867	T9578	T9575	prt	out,knocked
R5868	T9579	T9575	prep	in,knocked
R5869	T9580	T9581	compound	Gdf5,domains
R5870	T9581	T9579	pobj	domains,in
R5871	T9582	T9581	compound	expression,domains
R5872	T9583	T9571	punct	.,form
R5873	T9585	T9586	det	The,lack
R5874	T9586	T9587	nsubj	lack,be
R5875	T9588	T9586	prep	of,lack
R5876	T9589	T9590	compound	joint,fusions
R5877	T9590	T9588	pobj	fusions,of
R5878	T9591	T9586	prep	outside,lack
R5879	T9592	T9593	det	the,region
R5880	T9593	T9591	pobj	region,outside
R5881	T9594	T9593	compound	ankle,region
R5882	T9595	T9587	aux	could,be
R5883	T9596	T9587	prep	due,be
R5884	T9597	T9596	prep	to,due
R5885	T9598	T9597	pobj	differences,to
R5886	T9599	T9598	prep	in,differences
R5887	T9600	T9599	pobj	requirement,in
R5888	T9601	T9600	prep	for,requirement
R5889	T9602	T9603	compound	BMP,signaling
R5890	T9603	T9601	pobj	signaling,for
R5891	T9604	T9598	prep	in,differences
R5892	T9605	T9606	amod	different,joints
R5893	T9606	T9604	pobj	joints,in
R5894	T9607	T9597	punct	", ",to
R5895	T9608	T9597	conj	to,to
R5896	T9609	T9610	amod	compensating,expression
R5897	T9610	T9608	pobj	expression,to
R5898	T9611	T9610	prep	of,expression
R5899	T9612	T9613	amod	other,receptors
R5900	T9613	T9611	pobj	receptors,of
R5901	T9614	T9613	compound	BMP,receptors
R5902	T9615	T9613	prep	outside,receptors
R5903	T9616	T9617	det	the,ankles
R5904	T9617	T9615	pobj	ankles,outside
R5905	T9618	T9608	punct	", ",to
R5906	T9619	T9608	cc	or,to
R5907	T9620	T9608	conj	to,to
R5908	T9621	T9620	pobj	differences,to
R5909	T9622	T9621	prep	in,differences
R5910	T9623	T9624	det	the,timing
R5911	T9624	T9622	pobj	timing,in
R5912	T9625	T9624	amod	detailed,timing
R5913	T9626	T9624	prep	of,timing
R5914	T9627	T9628	compound	Gdf5,Cre
R5915	T9628	T9630	npadvmod	Cre,stimulated
R5916	T9629	T9628	punct	-,Cre
R5917	T9630	T9631	amod	stimulated,inactivation
R5918	T9631	T9626	pobj	inactivation,of
R5919	T9632	T9631	compound	gene,inactivation
R5920	T9633	T9631	prep	in,inactivation
R5921	T9634	T9633	pobj	ankles,in
R5922	T9635	T9634	cc	and,ankles
R5923	T9636	T9637	amod	other,regions
R5924	T9637	T9634	conj	regions,ankles
R5925	T9638	T9637	compound	joint,regions
R5926	T9639	T9587	punct	.,be
R5927	T9641	T9642	nsubj	Comparison,suggests
R5928	T9643	T9641	prep	of,Comparison
R5929	T9644	T9645	det	the,expression
R5930	T9645	T9643	pobj	expression,of
R5931	T9646	T9645	prep	of,expression
R5932	T9647	T9648	det	the,marker
R5933	T9648	T9646	pobj	marker,of
R5934	T9649	T9648	compound	HPLAP,marker
R5935	T9650	T9648	punct	(,marker
R5936	T9651	T9648	acl	driven,marker
R5937	T9652	T9651	advmod	directly,driven
R5938	T9653	T9651	agent	by,driven
R5939	T9654	T9655	compound	Gdf5,elements
R5940	T9655	T9653	pobj	elements,by
R5941	T9656	T9655	compound	control,elements
R5942	T9657	T9648	punct	),marker
R5943	T9658	T9648	cc	and,marker
R5944	T9659	T9660	det	the,marker
R5945	T9660	T9648	conj	marker,marker
R5946	T9661	T9662	compound	R26R,LACZ
R5947	T9662	T9660	compound	LACZ,marker
R5948	T9663	T9660	punct	(,marker
R5949	T9664	T9660	acl	expressed,marker
R5950	T9665	T9664	advcl	following,expressed
R5951	T9666	T9667	compound	Gdf5,Cre
R5952	T9667	T9669	compound	Cre,recombination
R5953	T9668	T9667	punct	-,Cre
R5954	T9669	T9665	dobj	recombination,following
R5955	T9670	T9660	punct	),marker
R5956	T9671	T9672	mark	that,delayed
R5957	T9672	T9642	ccomp	delayed,suggests
R5958	T9673	T9674	npadvmod	recombination,stimulated
R5959	T9674	T9676	amod	stimulated,changes
R5960	T9675	T9674	punct	-,stimulated
R5961	T9676	T9672	nsubjpass	changes,delayed
R5962	T9677	T9676	prep	in,changes
R5963	T9678	T9679	compound	gene,expression
R5964	T9679	T9677	pobj	expression,in
R5965	T9680	T9672	aux	may,delayed
R5966	T9681	T9672	auxpass	be,delayed
R5967	T9682	T9672	prep	for,delayed
R5968	T9683	T9684	det	a,d
R5969	T9684	T9682	pobj	d,for
R5970	T9685	T9686	quantmod	0.5,1
R5971	T9686	T9684	nummod	1,d
R5972	T9687	T9686	punct	–,1
R5973	T9688	T9672	prep	in,delayed
R5974	T9689	T9690	det	the,region
R5975	T9690	T9688	pobj	region,in
R5976	T9691	T9690	compound	digit,region
R5977	T9692	T9693	punct	(,see
R5978	T9693	T9642	parataxis	see,suggests
R5979	T9694	T9693	dobj	Figure,see
R5980	T9695	T9694	nummod	1C,Figure
R5981	T9696	T9693	punct	),see
R5982	T9697	T9642	punct	.,suggests
R5983	T9699	T9700	prep	In,persist
R5984	T9701	T9699	pobj	addition,In
R5985	T9702	T9700	punct	", ",persist
R5986	T9703	T9700	nsubj	levels,persist
R5987	T9704	T9703	prep	of,levels
R5988	T9705	T9706	compound	Bmpr1a,mRNA
R5989	T9706	T9704	pobj	mRNA,of
R5990	T9707	T9706	cc	and,mRNA
R5991	T9708	T9706	conj	protein,mRNA
R5992	T9709	T9700	aux	may,persist
R5993	T9710	T9700	prep	for,persist
R5994	T9711	T9712	det	some,time
R5995	T9712	T9710	pobj	time,for
R5996	T9713	T9700	prep	following,persist
R5997	T9714	T9715	compound	Gdf5,Cre
R5998	T9715	T9717	npadvmod	Cre,stimulated
R5999	T9716	T9715	punct	-,Cre
R6000	T9717	T9718	amod	stimulated,recombination
R6001	T9718	T9713	pobj	recombination,following
R6002	T9719	T9700	punct	", ",persist
R6003	T9720	T9700	advcl	making,persist
R6004	T9721	T9722	nsubj	it,possible
R6005	T9722	T9720	ccomp	possible,making
R6006	T9723	T9724	aux	to,bypass
R6007	T9724	T9722	advcl	bypass,possible
R6008	T9725	T9726	det	an,requirement
R6009	T9726	T9724	dobj	requirement,bypass
R6010	T9727	T9726	amod	early,requirement
R6011	T9728	T9726	prep	for,requirement
R6012	T9729	T9728	pobj	Bmpr1a,for
R6013	T9730	T9726	prep	in,requirement
R6014	T9731	T9732	compound	joint,formation
R6015	T9732	T9730	pobj	formation,in
R6016	T9733	T9732	prep	at,formation
R6017	T9734	T9735	det	some,locations
R6018	T9735	T9733	pobj	locations,at
R6019	T9736	T9700	punct	.,persist
R6020	T9738	T9739	prep	Following,result
R6021	T9740	T9741	det	the,decay
R6022	T9741	T9738	pobj	decay,Following
R6023	T9742	T9741	prep	of,decay
R6024	T9743	T9744	compound	Bmpr1a,mRNA
R6025	T9744	T9742	pobj	mRNA,of
R6026	T9745	T9744	cc	and,mRNA
R6027	T9746	T9744	conj	protein,mRNA
R6028	T9747	T9739	punct	", ",result
R6029	T9748	T9749	det	the,strategy
R6030	T9749	T9739	nsubj	strategy,result
R6031	T9750	T9751	compound	Gdf5,Cre
R6032	T9751	T9749	compound	Cre,strategy
R6033	T9752	T9751	punct	-,Cre
R6034	T9753	T9739	aux	should,result
R6035	T9754	T9739	prep	in,result
R6036	T9755	T9756	amod	permanent,inactivation
R6037	T9756	T9754	pobj	inactivation,in
R6038	T9757	T9756	prep	of,inactivation
R6039	T9758	T9759	compound	Bmpr1a,function
R6040	T9759	T9757	pobj	function,of
R6041	T9760	T9756	prep	in,inactivation
R6042	T9761	T9762	amod	recombined,cells
R6043	T9762	T9760	pobj	cells,in
R6044	T9763	T9739	punct	.,result
R6045	T9765	T9766	det	This,system
R6046	T9766	T9767	nsubj	system,provides
R6047	T9768	T9767	advmod	thus,provides
R6048	T9769	T9767	dobj	one,provides
R6049	T9770	T9769	prep	of,one
R6050	T9771	T9772	det	the,tests
R6051	T9772	T9770	pobj	tests,of
R6052	T9773	T9772	amod	first,tests
R6053	T9774	T9772	amod	strong,tests
R6054	T9775	T9772	amod	genetic,tests
R6055	T9776	T9772	prep	of,tests
R6056	T9777	T9778	compound	Bmpr1a,function
R6057	T9778	T9776	pobj	function,of
R6058	T9779	T9767	prep	at,provides
R6059	T9780	T9781	amod	later,stages
R6060	T9781	T9779	pobj	stages,at
R6061	T9782	T9781	prep	of,stages
R6062	T9783	T9784	compound	joint,development
R6063	T9784	T9782	pobj	development,of
R6064	T9785	T9767	punct	.,provides
R6065	T9787	T9788	prep	Despite,are
R6066	T9789	T9790	det	the,appearance
R6067	T9790	T9787	pobj	appearance,Despite
R6068	T9791	T9790	amod	normal,appearance
R6069	T9792	T9790	prep	of,appearance
R6070	T9793	T9794	amod	articular,regions
R6071	T9794	T9792	pobj	regions,of
R6072	T9795	T9794	cc	and,regions
R6073	T9796	T9797	compound	gene,expression
R6074	T9797	T9794	conj	expression,regions
R6075	T9798	T9799	advmod	immediately,after
R6076	T9799	T9790	prep	after,appearance
R6077	T9800	T9799	pobj	birth,after
R6078	T9801	T9788	punct	", ",are
R6079	T9802	T9803	npadvmod	Bmpr1a,deficient
R6080	T9803	T9805	amod	deficient,animals
R6081	T9804	T9803	punct	-,deficient
R6082	T9805	T9788	nsubj	animals,are
R6083	T9806	T9788	acomp	unable,are
R6084	T9807	T9808	aux	to,maintain
R6085	T9808	T9806	xcomp	maintain,unable
R6086	T9809	T9810	det	the,state
R6087	T9810	T9808	dobj	state,maintain
R6088	T9811	T9810	amod	normal,state
R6089	T9812	T9810	amod	differentiated,state
R6090	T9813	T9810	prep	of,state
R6091	T9814	T9815	amod	articular,cartilage
R6092	T9815	T9813	pobj	cartilage,of
R6093	T9816	T9817	mark	as,continue
R6094	T9817	T9808	advcl	continue,maintain
R6095	T9818	T9817	nsubj	they,continue
R6096	T9819	T9820	aux	to,develop
R6097	T9820	T9817	xcomp	develop,continue
R6098	T9821	T9820	cc	and,develop
R6099	T9822	T9820	conj	age,develop
R6100	T9823	T9788	punct	.,are
R6101	T9825	T9826	det	These,results
R6102	T9826	T9827	nsubj	results,suggest
R6103	T9828	T9829	mark	that,is
R6104	T9829	T9827	ccomp	is,suggest
R6105	T9830	T9831	compound	BMP,receptor
R6106	T9831	T9832	compound	receptor,signaling
R6107	T9832	T9829	nsubj	signaling,is
R6108	T9833	T9829	acomp	essential,is
R6109	T9834	T9833	prep	for,essential
R6110	T9835	T9836	amod	continued,health
R6111	T9836	T9834	pobj	health,for
R6112	T9837	T9836	cc	and,health
R6113	T9838	T9836	conj	integrity,health
R6114	T9839	T9838	prep	of,integrity
R6115	T9840	T9841	amod	articular,cartilage
R6116	T9841	T9839	pobj	cartilage,of
R6117	T9842	T9829	prep	in,is
R6118	T9843	T9844	det	the,period
R6119	T9844	T9842	pobj	period,in
R6120	T9845	T9844	amod	postnatal,period
R6121	T9846	T9827	punct	.,suggest
R6122	T9848	T9849	amod	Articular,cartilage
R6123	T9849	T9850	nsubj	cartilage,is
R6124	T9851	T9852	det	a,component
R6125	T9852	T9850	attr	component,is
R6126	T9853	T9852	amod	key,component
R6127	T9854	T9852	prep	of,component
R6128	T9855	T9856	amod	synovial,joints
R6129	T9856	T9854	pobj	joints,of
R6130	T9857	T9850	cc	and,is
R6131	T9858	T9850	conj	is,is
R6132	T9859	T9858	attr	one,is
R6133	T9860	T9859	prep	of,one
R6134	T9861	T9862	det	the,regions
R6135	T9862	T9860	pobj	regions,of
R6136	T9863	T9862	amod	few,regions
R6137	T9864	T9862	prep	in,regions
R6138	T9865	T9866	det	the,skeleton
R6139	T9866	T9864	pobj	skeleton,in
R6140	T9867	T9868	advmod	where,maintained
R6141	T9868	T9862	relcl	maintained,regions
R6142	T9869	T9868	nsubjpass	cartilage,maintained
R6143	T9870	T9868	auxpass	is,maintained
R6144	T9871	T9868	prep	into,maintained
R6145	T9872	T9871	pobj	adulthood,into
R6146	T9873	T9850	punct	.,is
R6147	T9875	T9876	prep	Despite,known
R6148	T9877	T9878	det	the,importance
R6149	T9878	T9875	pobj	importance,Despite
R6150	T9879	T9878	prep	of,importance
R6151	T9880	T9881	amod	articular,cartilage
R6152	T9881	T9879	pobj	cartilage,of
R6153	T9882	T9878	prep	in,importance
R6154	T9883	T9884	compound	joint,health
R6155	T9884	T9882	pobj	health,in
R6156	T9885	T9884	cc	and,health
R6157	T9886	T9884	conj	mobility,health
R6158	T9887	T9876	punct	", ",known
R6159	T9888	T9876	nsubjpass	little,known
R6160	T9889	T9876	auxpass	is,known
R6161	T9890	T9876	prep	about,known
R6162	T9891	T9892	det	the,factors
R6163	T9892	T9890	pobj	factors,about
R6164	T9893	T9894	dep	that,create
R6165	T9894	T9892	relcl	create,factors
R6166	T9895	T9894	cc	and,create
R6167	T9896	T9894	conj	maintain,create
R6168	T9897	T9896	dobj	it,maintain
R6169	T9898	T9896	prep	in,maintain
R6170	T9899	T9900	amod	thin,layers
R6171	T9900	T9898	pobj	layers,in
R6172	T9901	T9900	prep	at,layers
R6173	T9902	T9903	det	the,ends
R6174	T9903	T9901	pobj	ends,at
R6175	T9904	T9903	prep	of,ends
R6176	T9905	T9906	amod	long,bones
R6177	T9906	T9904	pobj	bones,of
R6178	T9907	T9876	punct	.,known
R6179	T9909	T9910	prep	In,retains
R6180	T9911	T9912	poss	our,experiments
R6181	T9912	T9909	pobj	experiments,In
R6182	T9913	T9910	punct	", ",retains
R6183	T9914	T9915	amod	articular,cartilage
R6184	T9915	T9916	npadvmod	cartilage,lacking
R6185	T9916	T9917	amod	lacking,Bmpr1a
R6186	T9917	T9910	nsubj	Bmpr1a,retains
R6187	T9918	T9919	det	some,characteristics
R6188	T9919	T9910	dobj	characteristics,retains
R6189	T9920	T9919	amod	normal,characteristics
R6190	T9921	T9910	punct	", ",retains
R6191	T9922	T9923	mark	in,maintains
R6192	T9923	T9910	advcl	maintains,retains
R6193	T9924	T9923	mark	that,maintains
R6194	T9925	T9923	nsubj	it,maintains
R6195	T9926	T9927	det	a,rate
R6196	T9927	T9923	dobj	rate,maintains
R6197	T9928	T9929	advmod	very,low
R6198	T9929	T9927	amod	low,rate
R6199	T9930	T9927	compound	proliferation,rate
R6200	T9931	T9923	punct	", ",maintains
R6201	T9932	T9933	aux	does,express
R6202	T9933	T9923	conj	express,maintains
R6203	T9934	T9933	neg	not,express
R6204	T9935	T9933	dobj	Col1a1,express
R6205	T9936	T9933	punct	", ",express
R6206	T9937	T9933	cc	and,express
R6207	T9938	T9933	conj	continues,express
R6208	T9939	T9940	aux	to,express
R6209	T9940	T9938	xcomp	express,continues
R6210	T9941	T9940	dobj	SOX9,express
R6211	T9942	T9941	punct	", ",SOX9
R6212	T9943	T9944	det	a,factor
R6213	T9944	T9941	appos	factor,SOX9
R6214	T9945	T9944	amod	major,factor
R6215	T9946	T9944	compound	transcription,factor
R6216	T9947	T9944	acl	regulating,factor
R6217	T9948	T9947	dobj	expression,regulating
R6218	T9949	T9948	prep	of,expression
R6219	T9950	T9951	amod	structural,components
R6220	T9951	T9949	pobj	components,of
R6221	T9952	T9951	prep	of,components
R6222	T9953	T9954	compound	cartilage,matrix
R6223	T9954	T9952	pobj	matrix,of
R6224	T9955	T9910	punct	.,retains
R6225	T9957	T9958	advmod	However,fail
R6226	T9959	T9958	punct	", ",fail
R6227	T9960	T9958	nsubj	several,fail
R6228	T9961	T9960	prep	of,several
R6229	T9962	T9963	det	the,components
R6230	T9963	T9961	pobj	components,of
R6231	T9964	T9965	advmod	most,prominent
R6232	T9965	T9963	amod	prominent,components
R6233	T9966	T9963	amod	structural,components
R6234	T9967	T9963	prep	of,components
R6235	T9968	T9969	compound	cartilage,matrix
R6236	T9969	T9967	pobj	matrix,of
R6237	T9970	T9971	aux	to,maintained
R6238	T9971	T9958	xcomp	maintained,fail
R6239	T9972	T9971	auxpass	be,maintained
R6240	T9973	T9971	prep	in,maintained
R6241	T9974	T9975	compound	mutant,animals
R6242	T9975	T9973	pobj	animals,in
R6243	T9976	T9971	punct	", ",maintained
R6244	T9977	T9971	advcl	resulting,maintained
R6245	T9978	T9977	prep	in,resulting
R6246	T9979	T9980	amod	decreased,synthesis
R6247	T9980	T9978	pobj	synthesis,in
R6248	T9981	T9980	prep	of,synthesis
R6249	T9982	T9981	pobj	Col2a1,of
R6250	T9983	T9982	punct	", ",Col2a1
R6251	T9984	T9982	conj	Agg,Col2a1
R6252	T9985	T9984	punct	", ",Agg
R6253	T9986	T9984	cc	and,Agg
R6254	T9987	T9984	conj	proteoglycans,Agg
R6255	T9988	T9958	punct	.,fail
R6256	T9990	T9991	advmod	Therefore,appears
R6257	T9992	T9991	punct	", ",appears
R6258	T9993	T9991	nsubj	BMPR1A,appears
R6259	T9994	T9995	aux	to,maintain
R6260	T9995	T9991	xcomp	maintain,appears
R6261	T9996	T9997	amod	articular,cartilage
R6262	T9997	T9995	dobj	cartilage,maintain
R6263	T9998	T9999	advmod	primarily,through
R6264	T9999	T9995	prep	through,maintain
R6265	T10000	T9999	pcomp	inducing,through
R6266	T10001	T10000	dobj	expression,inducing
R6267	T10002	T10001	prep	of,expression
R6268	T10003	T10004	amod	key,components
R6269	T10004	T10002	pobj	components,of
R6270	T10005	T10004	compound	ECM,components
R6271	T10006	T9991	punct	.,appears
R6272	T10008	T10009	nsubj	It,is
R6273	T10010	T10009	acomp	interesting,is
R6274	T10011	T10012	mark	that,continues
R6275	T10012	T10009	ccomp	continues,is
R6276	T10013	T10014	det	the,factor
R6277	T10014	T10012	nsubj	factor,continues
R6278	T10015	T10016	compound	SOX9,transcription
R6279	T10016	T10014	compound	transcription,factor
R6280	T10017	T10018	aux	to,expressed
R6281	T10018	T10012	xcomp	expressed,continues
R6282	T10019	T10018	auxpass	be,expressed
R6283	T10020	T10018	prep	in,expressed
R6284	T10021	T10022	compound	mutant,cartilage
R6285	T10022	T10020	pobj	cartilage,in
R6286	T10023	T10018	prep	despite,expressed
R6287	T10024	T10023	pobj	loss,despite
R6288	T10025	T10024	prep	of,loss
R6289	T10026	T10025	pobj	Col2a1,of
R6290	T10027	T10026	punct	", ",Col2a1
R6291	T10028	T10029	det	a,target
R6292	T10029	T10026	appos	target,Col2a1
R6293	T10030	T10029	amod	direct,target
R6294	T10031	T10029	prep	of,target
R6295	T10032	T10033	det	this,factor
R6296	T10033	T10031	pobj	factor,of
R6297	T10034	T10033	compound	transcription,factor
R6298	T10035	T10036	punct	(,Bell
R6299	T10036	T10009	meta	Bell,is
R6300	T10037	T10036	nmod	et,Bell
R6301	T10038	T10036	nmod	al.,Bell
R6302	T10039	T10036	nummod	1997,Bell
R6303	T10040	T10036	punct	;,Bell
R6304	T10041	T10036	nmod	Lefebvre,Bell
R6305	T10042	T10036	nmod	et,Bell
R6306	T10043	T10036	nmod	al.,Bell
R6307	T10044	T10036	nummod	1997,Bell
R6308	T10045	T10036	punct	),Bell
R6309	T10046	T10009	punct	.,is
R6310	T10048	T10049	amod	Previous,studies
R6311	T10049	T10050	nsubj	studies,suggest
R6312	T10051	T10052	mark	that,modified
R6313	T10052	T10050	ccomp	modified,suggest
R6314	T10053	T10054	compound	SOX9,activity
R6315	T10054	T10052	nsubjpass	activity,modified
R6316	T10055	T10052	aux	can,modified
R6317	T10056	T10052	auxpass	be,modified
R6318	T10057	T10052	prep	by,modified
R6319	T10058	T10059	compound	protein,A
R6320	T10059	T10061	npadvmod	A,dependent
R6321	T10060	T10059	compound	kinase,A
R6322	T10061	T10066	amod	dependent,phosphorylation
R6323	T10062	T10059	punct	(,A
R6324	T10063	T10059	appos	PKA,A
R6325	T10064	T10059	punct	),A
R6326	T10065	T10061	punct	-,dependent
R6327	T10066	T10057	pobj	phosphorylation,by
R6328	T10067	T10066	compound	protein,phosphorylation
R6329	T10068	T10057	punct	", ",by
R6330	T10069	T10057	cc	or,by
R6331	T10070	T10057	conj	by,by
R6332	T10071	T10070	pobj	coexpression,by
R6333	T10072	T10071	prep	of,coexpression
R6334	T10073	T10074	nummod	two,proteins
R6335	T10074	T10072	pobj	proteins,of
R6336	T10075	T10074	amod	related,proteins
R6337	T10076	T10074	punct	", ",proteins
R6338	T10077	T10078	compound	L,SOX5
R6339	T10078	T10074	appos	SOX5,proteins
R6340	T10079	T10078	punct	-,SOX5
R6341	T10080	T10078	cc	and,SOX5
R6342	T10081	T10078	conj	SOX6,SOX5
R6343	T10082	T10083	punct	(,Lefebvre
R6344	T10083	T10050	meta	Lefebvre,suggest
R6345	T10084	T10083	nmod	et,Lefebvre
R6346	T10085	T10083	nmod	al.,Lefebvre
R6347	T10086	T10083	nummod	1998,Lefebvre
R6348	T10087	T10083	punct	;,Lefebvre
R6349	T10088	T10083	nmod	Huang,Lefebvre
R6350	T10089	T10083	nmod	et,Lefebvre
R6351	T10090	T10083	nmod	al.,Lefebvre
R6352	T10091	T10083	nummod	2000,Lefebvre
R6353	T10092	T10083	punct	),Lefebvre
R6354	T10093	T10050	punct	.,suggest
R6355	T10095	T10096	prep	In,reveals
R6356	T10097	T10095	pobj	addition,In
R6357	T10098	T10096	punct	", ",reveals
R6358	T10099	T10100	amod	close,examination
R6359	T10100	T10096	nsubj	examination,reveals
R6360	T10101	T10100	prep	of,examination
R6361	T10102	T10103	det	the,order
R6362	T10103	T10101	pobj	order,of
R6363	T10104	T10103	prep	of,order
R6364	T10105	T10104	pobj	genes,of
R6365	T10106	T10105	acl	induced,genes
R6366	T10107	T10106	prep	during,induced
R6367	T10108	T10109	compound	chicken,formation
R6368	T10109	T10107	pobj	formation,during
R6369	T10110	T10109	compound	digit,formation
R6370	T10111	T10112	mark	that,turns
R6371	T10112	T10096	ccomp	turns,reveals
R6372	T10113	T10112	nsubj	Sox9,turns
R6373	T10114	T10112	prt	on,turns
R6374	T10115	T10112	advmod	first,turns
R6375	T10116	T10112	punct	", ",turns
R6376	T10117	T10112	advcl	followed,turns
R6377	T10118	T10117	agent	by,followed
R6378	T10119	T10118	pobj	Bmpr1b,by
R6379	T10120	T10119	prep	with,Bmpr1b
R6380	T10121	T10122	compound	L,Sox5
R6381	T10122	T10120	pobj	Sox5,with
R6382	T10123	T10122	punct	-,Sox5
R6383	T10124	T10119	punct	", ",Bmpr1b
R6384	T10125	T10119	cc	and,Bmpr1b
R6385	T10126	T10127	advmod	then,Sox6
R6386	T10127	T10119	conj	Sox6,Bmpr1b
R6387	T10128	T10127	cc	and,Sox6
R6388	T10129	T10130	det	the,components
R6389	T10130	T10127	conj	components,Sox6
R6390	T10131	T10132	nmod	cartilage,matrix
R6391	T10132	T10130	nmod	matrix,components
R6392	T10133	T10130	amod	structural,components
R6393	T10134	T10130	appos	Col2a1,components
R6394	T10135	T10134	cc	and,Col2a1
R6395	T10136	T10134	conj	Agg,Col2a1
R6396	T10137	T10138	punct	(,Chimal
R6397	T10138	T10096	meta	Chimal,reveals
R6398	T10139	T10138	punct	-,Chimal
R6399	T10140	T10138	nmod	Monroy,Chimal
R6400	T10141	T10138	nmod	et,Chimal
R6401	T10142	T10138	nmod	al.,Chimal
R6402	T10143	T10138	nummod	2003,Chimal
R6403	T10144	T10138	punct	),Chimal
R6404	T10145	T10096	punct	.,reveals
R6405	T10147	T10148	det	These,results
R6406	T10148	T10149	nsubj	results,suggest
R6407	T10150	T10148	punct	", ",results
R6408	T10151	T10148	advmod	together,results
R6409	T10152	T10151	prep	with,together
R6410	T10153	T10154	det	the,pattern
R6411	T10154	T10152	pobj	pattern,with
R6412	T10155	T10154	amod	altered,pattern
R6413	T10156	T10154	prep	of,pattern
R6414	T10157	T10158	compound	gene,expression
R6415	T10158	T10156	pobj	expression,of
R6416	T10159	T10154	acl	seen,pattern
R6417	T10160	T10159	prep	in,seen
R6418	T10161	T10162	poss	our,mice
R6419	T10162	T10160	pobj	mice,in
R6420	T10163	T10164	npadvmod	Bmpr1a,deficient
R6421	T10164	T10162	amod	deficient,mice
R6422	T10165	T10164	punct	-,deficient
R6423	T10166	T10149	punct	", ",suggest
R6424	T10167	T10168	mark	that,act
R6425	T10168	T10149	ccomp	act,suggest
R6426	T10169	T10170	compound	BMPR1A,signaling
R6427	T10170	T10168	nsubj	signaling,act
R6428	T10171	T10168	aux	may,act
R6429	T10172	T10168	advmod	normally,act
R6430	T10173	T10174	aux	to,stimulate
R6431	T10174	T10168	xcomp	stimulate,act
R6432	T10175	T10174	dobj	SOX9,stimulate
R6433	T10176	T10174	prep	by,stimulate
R6434	T10177	T10178	amod	post-translational,modification
R6435	T10178	T10176	pobj	modification,by
R6436	T10179	T10178	compound	protein,modification
R6437	T10180	T10174	punct	", ",stimulate
R6438	T10181	T10174	cc	or,stimulate
R6439	T10182	T10183	aux	to,induce
R6440	T10183	T10174	conj	induce,stimulate
R6441	T10184	T10185	compound	L,Sox5
R6442	T10185	T10183	dobj	Sox5,induce
R6443	T10186	T10185	punct	-,Sox5
R6444	T10187	T10185	cc	or,Sox5
R6445	T10188	T10185	conj	Sox6,Sox5
R6446	T10189	T10183	prep	in,induce
R6447	T10190	T10189	pobj	cartilage,in
R6448	T10191	T10192	aux	to,maintain
R6449	T10192	T10183	advcl	maintain,induce
R6450	T10193	T10192	dobj	expression,maintain
R6451	T10194	T10193	prep	of,expression
R6452	T10195	T10196	compound	ECM,components
R6453	T10196	T10194	pobj	components,of
R6454	T10197	T10149	punct	.,suggest
R6455	T10199	T10200	det	These,models
R6456	T10200	T10201	nsubj	models,are
R6457	T10202	T10201	acomp	consistent,are
R6458	T10203	T10202	prep	with,consistent
R6459	T10204	T10205	det	the,ability
R6460	T10205	T10203	pobj	ability,with
R6461	T10206	T10205	prep	of,ability
R6462	T10207	T10206	pobj	BMP2,of
R6463	T10208	T10209	aux	to,increase
R6464	T10209	T10205	acl	increase,ability
R6465	T10210	T10209	preconj	both,increase
R6466	T10211	T10212	compound	PKA,activity
R6467	T10212	T10209	dobj	activity,increase
R6468	T10213	T10209	cc	and,increase
R6469	T10214	T10209	conj	induce,increase
R6470	T10215	T10214	dobj	expression,induce
R6471	T10216	T10215	prep	of,expression
R6472	T10217	T10216	pobj	Sox6,of
R6473	T10218	T10214	prep	in,induce
R6474	T10219	T10220	compound	tissue,cells
R6475	T10220	T10218	pobj	cells,in
R6476	T10221	T10220	compound	culture,cells
R6477	T10222	T10223	punct	(,Lee
R6478	T10223	T10201	meta	Lee,are
R6479	T10224	T10223	cc	and,Lee
R6480	T10225	T10223	conj	Chuong,Lee
R6481	T10226	T10225	nummod	1997,Chuong
R6482	T10227	T10225	punct	;,Chuong
R6483	T10228	T10225	conj	Fernandez,Chuong
R6484	T10229	T10228	punct	-,Fernandez
R6485	T10230	T10228	nmod	Lloris,Fernandez
R6486	T10231	T10228	nmod	et,Fernandez
R6487	T10232	T10228	nmod	al.,Fernandez
R6488	T10233	T10228	nummod	2003,Fernandez
R6489	T10234	T10228	punct	),Fernandez
R6490	T10235	T10201	punct	.,are
R6491	T10237	T10238	mark	Although,tried
R6492	T10238	T10241	advcl	tried,been
R6493	T10239	T10238	nsubj	we,tried
R6494	T10240	T10238	aux	have,tried
R6495	T10242	T10243	aux	to,monitor
R6496	T10243	T10238	xcomp	monitor,tried
R6497	T10244	T10245	det	the,expression
R6498	T10245	T10243	dobj	expression,monitor
R6499	T10246	T10245	prep	of,expression
R6500	T10247	T10248	compound	L,Sox5
R6501	T10248	T10246	pobj	Sox5,of
R6502	T10249	T10248	punct	-,Sox5
R6503	T10250	T10248	cc	or,Sox5
R6504	T10251	T10248	conj	Sox6,Sox5
R6505	T10252	T10243	prep	in,monitor
R6506	T10253	T10254	amod	postnatal,cartilage
R6507	T10254	T10252	pobj	cartilage,in
R6508	T10255	T10254	amod	articular,cartilage
R6509	T10256	T10243	punct	", ",monitor
R6510	T10257	T10243	cc	and,monitor
R6511	T10258	T10243	conj	test,monitor
R6512	T10259	T10260	det	the,state
R6513	T10260	T10258	dobj	state,test
R6514	T10261	T10260	compound	phosphorylation,state
R6515	T10262	T10260	prep	of,state
R6516	T10263	T10262	pobj	SOX9,of
R6517	T10264	T10258	advcl	using,test
R6518	T10265	T10266	advmod	previously,described
R6519	T10266	T10267	amod	described,reagents
R6520	T10267	T10264	dobj	reagents,using
R6521	T10268	T10269	punct	(,Lefebvre
R6522	T10269	T10258	meta	Lefebvre,test
R6523	T10270	T10269	nmod	et,Lefebvre
R6524	T10271	T10269	nmod	al.,Lefebvre
R6525	T10272	T10269	nummod	1998,Lefebvre
R6526	T10273	T10269	punct	;,Lefebvre
R6527	T10274	T10269	nmod	Huang,Lefebvre
R6528	T10275	T10269	nmod	et,Lefebvre
R6529	T10276	T10269	nmod	al.,Lefebvre
R6530	T10277	T10269	nummod	2000,Lefebvre
R6531	T10278	T10269	punct	),Lefebvre
R6532	T10279	T10241	punct	", ",been
R6533	T10280	T10241	nsubj	we,been
R6534	T10281	T10241	aux	have,been
R6535	T10282	T10241	acomp	unable,been
R6536	T10283	T10284	aux	to,obtain
R6537	T10284	T10282	xcomp	obtain,unable
R6538	T10285	T10286	amod	specific,signal
R6539	T10286	T10284	dobj	signal,obtain
R6540	T10287	T10284	prep	at,obtain
R6541	T10288	T10289	det	the,stages
R6542	T10289	T10287	pobj	stages,at
R6543	T10290	T10289	amod	late,stages
R6544	T10291	T10289	amod	postnatal,stages
R6545	T10292	T10289	acl	required,stages
R6546	T10293	T10294	punct	(,data
R6547	T10294	T10241	meta	data,been
R6548	T10295	T10294	amod	unpublished,data
R6549	T10296	T10294	punct	),data
R6550	T10297	T10241	punct	.,been
R6551	T10299	T10300	advmod	Furthermore,cause
R6552	T10301	T10300	punct	", ",cause
R6553	T10302	T10303	amod	null,mutations
R6554	T10303	T10300	nsubj	mutations,cause
R6555	T10304	T10303	prep	in,mutations
R6556	T10305	T10306	compound	L,Sox5
R6557	T10306	T10304	pobj	Sox5,in
R6558	T10307	T10306	punct	-,Sox5
R6559	T10308	T10306	cc	or,Sox5
R6560	T10309	T10306	conj	Sox,Sox5
R6561	T10310	T10309	punct	-,Sox
R6562	T10311	T10309	nummod	6,Sox
R6563	T10312	T10300	dobj	lethality,cause
R6564	T10313	T10312	prep	at,lethality
R6565	T10314	T10313	cc	or,at
R6566	T10315	T10316	advmod	soon,after
R6567	T10316	T10313	conj	after,at
R6568	T10317	T10316	pobj	birth,after
R6569	T10318	T10300	punct	", ",cause
R6570	T10319	T10300	cc	and,cause
R6571	T10320	T10321	det	no,effect
R6572	T10321	T10322	nsubjpass	effect,reported
R6573	T10322	T10300	conj	reported,cause
R6574	T10323	T10321	prep	on,effect
R6575	T10324	T10325	compound	cartilage,maintenance
R6576	T10325	T10323	pobj	maintenance,on
R6577	T10326	T10322	aux	has,reported
R6578	T10327	T10322	auxpass	been,reported
R6579	T10328	T10329	punct	(,Smits
R6580	T10329	T10322	meta	Smits,reported
R6581	T10330	T10329	nmod	et,Smits
R6582	T10331	T10329	nmod	al.,Smits
R6583	T10332	T10329	nummod	2001,Smits
R6584	T10333	T10329	punct	),Smits
R6585	T10334	T10322	punct	.,reported
R6586	T10336	T10337	advmod	However,seems
R6587	T10338	T10337	punct	", ",seems
R6588	T10339	T10337	nsubj	it,seems
R6589	T10340	T10337	oprd	likely,seems
R6590	T10341	T10342	mark	that,cooperate
R6591	T10342	T10337	ccomp	cooperate,seems
R6592	T10343	T10342	nsubj	these,cooperate
R6593	T10344	T10343	cc	or,these
R6594	T10345	T10346	amod	other,processes
R6595	T10346	T10343	conj	processes,these
R6596	T10347	T10346	acl	regulated,processes
R6597	T10348	T10347	agent	by,regulated
R6598	T10349	T10350	compound	BMP,signaling
R6599	T10350	T10348	pobj	signaling,by
R6600	T10351	T10342	prep	with,cooperate
R6601	T10352	T10351	pobj	SOX9,with
R6602	T10353	T10354	aux	to,induce
R6603	T10354	T10342	advcl	induce,cooperate
R6604	T10355	T10356	amod	target,genes
R6605	T10356	T10354	dobj	genes,induce
R6606	T10357	T10354	prep	in,induce
R6607	T10358	T10359	amod	articular,cartilage
R6608	T10359	T10357	pobj	cartilage,in
R6609	T10360	T10337	punct	.,seems
R6610	T10362	T10363	nsubj	Mutation,disrupts
R6611	T10364	T10362	prep	of,Mutation
R6612	T10365	T10364	pobj	Smad3,of
R6613	T10366	T10362	cc	or,Mutation
R6614	T10367	T10362	conj	expression,Mutation
R6615	T10368	T10367	prep	of,expression
R6616	T10369	T10370	amod	dominant,receptor
R6617	T10370	T10368	pobj	receptor,of
R6618	T10371	T10370	amod	negative,receptor
R6619	T10372	T10373	amod	transforming,β
R6620	T10373	T10370	nmod	β,receptor
R6621	T10374	T10373	nmod	growth,β
R6622	T10375	T10373	nmod	factor,β
R6623	T10376	T10373	punct	(,β
R6624	T10377	T10378	compound	TGF,β
R6625	T10378	T10373	appos	β,β
R6626	T10379	T10378	punct	-,β
R6627	T10380	T10373	punct	),β
R6628	T10381	T10370	nmod	type,receptor
R6629	T10382	T10381	nummod	II,type
R6630	T10383	T10363	advmod	also,disrupts
R6631	T10384	T10385	amod	normal,maintenance
R6632	T10385	T10363	dobj	maintenance,disrupts
R6633	T10386	T10387	amod	articular,cartilage
R6634	T10387	T10385	compound	cartilage,maintenance
R6635	T10388	T10389	punct	(,Serra
R6636	T10389	T10363	meta	Serra,disrupts
R6637	T10390	T10389	nmod	et,Serra
R6638	T10391	T10389	nmod	al.,Serra
R6639	T10392	T10389	nummod	1997,Serra
R6640	T10393	T10389	punct	;,Serra
R6641	T10394	T10389	nmod	Yang,Serra
R6642	T10395	T10389	nmod	et,Serra
R6643	T10396	T10389	nmod	al.,Serra
R6644	T10397	T10389	nummod	2001,Serra
R6645	T10398	T10389	punct	),Serra
R6646	T10399	T10363	punct	.,disrupts
R6647	T10401	T10402	det	Both,manipulations
R6648	T10402	T10403	nsubj	manipulations,disrupt
R6649	T10404	T10403	aux	should,disrupt
R6650	T10405	T10403	dobj	TGFβ,disrupt
R6651	T10406	T10407	amod	rather,than
R6652	T10407	T10405	prep	than,TGFβ
R6653	T10408	T10409	compound	BMP,signaling
R6654	T10409	T10407	pobj	signaling,than
R6655	T10410	T10403	punct	", ",disrupt
R6656	T10411	T10403	cc	and,disrupt
R6657	T10412	T10413	det	both,manipulations
R6658	T10413	T10414	nsubj	manipulations,cause
R6659	T10414	T10403	conj	cause,disrupt
R6660	T10415	T10416	amod	articular,cartilage
R6661	T10416	T10417	nsubj	cartilage,hypertrophy
R6662	T10417	T10414	ccomp	hypertrophy,cause
R6663	T10418	T10417	aux	to,hypertrophy
R6664	T10419	T10417	cc	and,hypertrophy
R6665	T10420	T10421	auxpass	be,replaced
R6666	T10421	T10417	conj	replaced,hypertrophy
R6667	T10422	T10421	agent	by,replaced
R6668	T10423	T10422	pobj	bone,by
R6669	T10424	T10414	punct	.,cause
R6670	T10426	T10427	prep	In,showed
R6671	T10428	T10426	pobj	contrast,In
R6672	T10429	T10427	punct	", ",showed
R6673	T10430	T10431	poss	our,analysis
R6674	T10431	T10427	nsubj	analysis,showed
R6675	T10432	T10431	prep	of,analysis
R6676	T10433	T10434	nmod	Bmpr1a,mutant
R6677	T10434	T10435	nmod	mutant,cartilage
R6678	T10435	T10432	pobj	cartilage,of
R6679	T10436	T10435	amod	articular,cartilage
R6680	T10437	T10438	det	a,loss
R6681	T10438	T10427	dobj	loss,showed
R6682	T10439	T10438	prep	of,loss
R6683	T10440	T10441	compound	ECM,components
R6684	T10441	T10439	pobj	components,of
R6685	T10442	T10438	punct	", ",loss
R6686	T10443	T10438	cc	but,loss
R6687	T10444	T10445	det	no,signs
R6688	T10445	T10438	conj	signs,loss
R6689	T10446	T10445	prep	of,signs
R6690	T10447	T10446	pobj	hypertrophy,of
R6691	T10448	T10447	cc	or,hypertrophy
R6692	T10449	T10450	compound	bone,replacement
R6693	T10450	T10447	conj	replacement,hypertrophy
R6694	T10451	T10427	punct	.,showed
R6695	T10453	T10454	advmod	Therefore,playing
R6696	T10455	T10454	punct	", ",playing
R6697	T10456	T10457	nmod	TGFβ,signaling
R6698	T10457	T10454	nsubj	signaling,playing
R6699	T10458	T10456	cc	and,TGFβ
R6700	T10459	T10456	conj	BMP,TGFβ
R6701	T10460	T10454	aux	are,playing
R6702	T10461	T10462	amod	distinct,roles
R6703	T10462	T10454	dobj	roles,playing
R6704	T10463	T10461	cc	but,distinct
R6705	T10464	T10461	conj	necessary,distinct
R6706	T10465	T10466	aux	to,maintain
R6707	T10466	T10462	advcl	maintain,roles
R6708	T10467	T10468	amod	articular,cartilage
R6709	T10468	T10466	dobj	cartilage,maintain
R6710	T10469	T10454	punct	.,playing
R6711	T10471	T10472	mark	Although,isolated
R6712	T10472	T10476	advcl	isolated,stimulated
R6713	T10473	T10472	nsubjpass	BMPs,isolated
R6714	T10474	T10472	auxpass	were,isolated
R6715	T10475	T10472	advmod	originally,isolated
R6716	T10477	T10472	prep	on,isolated
R6717	T10478	T10479	det	the,basis
R6718	T10479	T10477	pobj	basis,on
R6719	T10480	T10479	prep	of,basis
R6720	T10481	T10482	poss	their,ability
R6721	T10482	T10480	pobj	ability,of
R6722	T10483	T10484	aux	to,induce
R6723	T10484	T10482	acl	induce,ability
R6724	T10485	T10486	amod	ectopic,formation
R6725	T10486	T10484	dobj	formation,induce
R6726	T10487	T10486	compound	bone,formation
R6727	T10488	T10476	punct	", ",stimulated
R6728	T10489	T10490	poss	their,presence
R6729	T10490	T10476	nsubj	presence,stimulated
R6730	T10521	T10520	nmod	et,Erlacher
R6731	T10491	T10490	prep	in,presence
R6732	T10522	T10520	nmod	al.,Erlacher
R6733	T10492	T10493	amod	articular,cartilage
R6734	T10523	T10520	nummod	1998,Erlacher
R6735	T10493	T10491	pobj	cartilage,in
R6736	T10494	T10490	cc	and,presence
R6737	T10524	T10520	punct	;,Erlacher
R6738	T10495	T10496	amod	strong,effect
R6739	T10525	T10520	conj	Edwards,Erlacher
R6740	T10526	T10525	cc	and,Edwards
R6741	T10496	T10490	conj	effect,presence
R6742	T10527	T10525	conj	Francis,Edwards
R6743	T10528	T10527	punct	-,Francis
R6744	T10497	T10496	prep	on,effect
R6745	T10529	T10527	nmod	West,Francis
R6746	T10530	T10527	nummod	2001,Francis
R6747	T10531	T10527	punct	;,Francis
R6748	T10498	T10499	compound	cartilage,formation
R6749	T10532	T10527	conj	Chubinskaya,Francis
R6750	T10533	T10532	cc	and,Chubinskaya
R6751	T10499	T10497	pobj	formation,on
R6752	T10534	T10532	conj	Kuettner,Chubinskaya
R6753	T10535	T10534	nummod	2003,Kuettner
R6754	T10500	T10476	aux	has,stimulated
R6755	T10536	T10534	punct	),Kuettner
R6756	T10537	T10476	punct	.,stimulated
R6757	T10539	T10540	det	The,failure
R6758	T10540	T10541	nsubj	failure,suggests
R6759	T10501	T10476	dobj	interest,stimulated
R6760	T10542	T10543	aux	to,maintain
R6761	T10543	T10540	acl	maintain,failure
R6762	T10502	T10501	prep	in,interest
R6763	T10544	T10545	amod	articular,cartilage
R6764	T10545	T10543	dobj	cartilage,maintain
R6765	T10546	T10543	prep	in,maintain
R6766	T10503	T10502	pcomp	using,in
R6767	T10547	T10548	det	the,absence
R6768	T10548	T10546	pobj	absence,in
R6769	T10549	T10548	prep	of,absence
R6770	T10504	T10503	dobj	them,using
R6771	T10550	T10551	amod	normal,function
R6772	T10551	T10549	pobj	function,of
R6773	T10505	T10506	aux	to,repair
R6774	T10552	T10551	compound	BMPR1A,function
R6775	T10506	T10503	advcl	repair,using
R6776	T10553	T10554	mark	that,be
R6777	T10554	T10541	ccomp	be,suggests
R6778	T10555	T10554	nsubj	ligands,be
R6779	T10556	T10555	cc	or,ligands
R6780	T10557	T10558	amod	small,agonists
R6781	T10507	T10506	cc	or,repair
R6782	T10558	T10555	conj	agonists,ligands
R6783	T10559	T10558	compound	molecule,agonists
R6784	T10560	T10561	dep	that,interact
R6785	T10508	T10506	conj	regenerate,repair
R6786	T10561	T10555	relcl	interact,ligands
R6787	T10562	T10561	advmod	specifically,interact
R6788	T10563	T10561	prep	with,interact
R6789	T10509	T10510	compound	cartilage,defects
R6790	T10564	T10565	det	this,subtype
R6791	T10565	T10563	pobj	subtype,with
R6792	T10566	T10565	compound	receptor,subtype
R6793	T10510	T10508	dobj	defects,regenerate
R6794	T10567	T10554	aux	may,be
R6795	T10568	T10569	advmod	particularly,good
R6796	T10569	T10570	amod	good,candidates
R6797	T10511	T10508	prep	in,regenerate
R6798	T10570	T10554	attr	candidates,be
R6799	T10571	T10570	prep	for,candidates
R6800	T10512	T10513	compound	adult,animals
R6801	T10572	T10571	pcomp	designing,for
R6802	T10573	T10574	amod	new,approaches
R6803	T10574	T10572	dobj	approaches,designing
R6804	T10513	T10511	pobj	animals,in
R6805	T10575	T10576	aux	to,maintain
R6806	T10576	T10574	acl	maintain,approaches
R6807	T10514	T10515	punct	(,Chang
R6808	T10577	T10576	cc	or,maintain
R6809	T10578	T10576	conj	heal,maintain
R6810	T10579	T10580	amod	articular,cartilage
R6811	T10580	T10578	dobj	cartilage,heal
R6812	T10581	T10578	prep	at,heal
R6813	T10582	T10583	amod	postnatal,stages
R6814	T10515	T10476	meta	Chang,stimulated
R6815	T10583	T10581	pobj	stages,at
R6816	T10584	T10541	punct	.,suggests
R6817	T10516	T10515	nmod	et,Chang
R6818	T10586	T10587	nsubj	Lack,results
R6819	T10517	T10515	nmod	al.,Chang
R6820	T10588	T10586	prep	of,Lack
R6821	T10589	T10590	compound	Bmpr1a,function
R6822	T10590	T10588	pobj	function,of
R6823	T10518	T10515	nummod	1994,Chang
R6824	T10591	T10590	prep	in,function
R6825	T10592	T10593	amod	articular,cartilage
R6826	T10519	T10515	punct	;,Chang
R6827	T10593	T10591	pobj	cartilage,in
R6828	T10520	T10515	conj	Erlacher,Chang
R6829	T10594	T10587	prep	in,results
R6830	T10595	T10596	amod	severe,fibrillation
R6831	T10596	T10594	pobj	fibrillation,in
R6832	T10597	T10596	prep	of,fibrillation
R6833	T10627	T10625	pobj	signaling,in
R6834	T10598	T10599	det	the,surface
R6835	T10628	T10623	advmod	also,contribute
R6836	T10599	T10597	pobj	surface,of
R6837	T10600	T10599	amod	articular,surface
R6838	T10601	T10596	cc	and,fibrillation
R6839	T10629	T10623	prep	to,contribute
R6840	T10602	T10596	conj	loss,fibrillation
R6841	T10603	T10602	prep	of,loss
R6842	T10630	T10631	amod	human,disease
R6843	T10604	T10605	compound	joint,mobility
R6844	T10605	T10603	pobj	mobility,of
R6845	T10606	T10587	punct	.,results
R6846	T10631	T10629	pobj	disease,to
R6847	T10608	T10609	det	The,development
R6848	T10609	T10610	nsubj	development,raises
R6849	T10632	T10631	compound	joint,disease
R6850	T10633	T10610	punct	.,raises
R6851	T10611	T10609	prep	of,development
R6852	T10635	T10636	nsubjpass	Osteoarthritis,known
R6853	T10612	T10613	amod	severe,symptoms
R6854	T10613	T10611	pobj	symptoms,of
R6855	T10614	T10613	compound	arthritis,symptoms
R6856	T10615	T10609	prep	in,development
R6857	T10637	T10636	auxpass	is,known
R6858	T10616	T10617	npadvmod	Bmpr1a,deficient
R6859	T10617	T10619	amod	deficient,mice
R6860	T10618	T10617	punct	-,deficient
R6861	T10619	T10615	pobj	mice,in
R6862	T10620	T10621	det	the,possibility
R6863	T10621	T10610	dobj	possibility,raises
R6864	T10638	T10639	aux	to,have
R6865	T10639	T10636	xcomp	have,known
R6866	T10622	T10623	mark	that,contribute
R6867	T10640	T10641	det	a,component
R6868	T10623	T10621	acl	contribute,possibility
R6869	T10624	T10623	nsubj	defects,contribute
R6870	T10641	T10639	dobj	component,have
R6871	T10625	T10624	prep	in,defects
R6872	T10626	T10627	compound	BMP,signaling
R6873	T10642	T10641	amod	significant,component
R6874	T10643	T10641	amod	genetic,component
R6875	T10644	T10636	punct	", ",known
R6876	T10645	T10636	cc	but,known
R6877	T10733	T10731	pobj	study,in
R6878	T10646	T10647	nsubj	it,involves
R6879	T10734	T10725	acomp	heterozygous,were
R6880	T10735	T10734	prep	for,heterozygous
R6881	T10647	T10636	conj	involves,known
R6882	T10736	T10737	det	a,allele
R6883	T10737	T10735	pobj	allele,for
R6884	T10738	T10737	amod	null,allele
R6885	T10648	T10647	advmod	likely,involves
R6886	T10739	T10737	prep	of,allele
R6887	T10740	T10739	pobj	Bmpr1a,of
R6888	T10741	T10725	punct	", ",were
R6889	T10649	T10650	amod	multiple,factors
R6890	T10742	T10725	cc	and,were
R6891	T10743	T10744	nsubj	they,showed
R6892	T10744	T10725	conj	showed,were
R6893	T10650	T10647	dobj	factors,involves
R6894	T10745	T10746	amod	little,sign
R6895	T10746	T10744	dobj	sign,showed
R6896	T10651	T10650	amod	genetic,factors
R6897	T10747	T10746	prep	of,sign
R6898	T10748	T10747	pobj	osteoarthritis,of
R6899	T10749	T10750	advmod	even,late
R6900	T10652	T10653	dep	that,been
R6901	T10750	T10744	advmod	late,showed
R6902	T10751	T10750	prep	in,late
R6903	T10752	T10751	pobj	life,in
R6904	T10653	T10650	relcl	been,factors
R6905	T10753	T10744	punct	.,showed
R6906	T10654	T10653	aux	have,been
R6907	T10755	T10756	amod	Several,regions
R6908	T10756	T10758	nsubjpass	regions,linked
R6909	T10757	T10756	compound	chromosome,regions
R6910	T10655	T10653	acomp	difficult,been
R6911	T10759	T10758	aux	have,linked
R6912	T10760	T10758	auxpass	been,linked
R6913	T10761	T10758	advmod	previously,linked
R6914	T10762	T10758	prep	to,linked
R6915	T10763	T10764	compound	arthritis,phenotypes
R6916	T10656	T10657	aux	to,identify
R6917	T10764	T10762	pobj	phenotypes,to
R6918	T10765	T10758	prep	in,linked
R6919	T10766	T10765	pobj	humans,in
R6920	T10657	T10655	advcl	identify,difficult
R6921	T10767	T10758	advcl	using,linked
R6922	T10768	T10769	preconj	either,studies
R6923	T10769	T10767	dobj	studies,using
R6924	T10658	T10659	punct	(,Spector
R6925	T10770	T10769	compound	association,studies
R6926	T10771	T10769	prep	in,studies
R6927	T10772	T10771	pobj	populations,in
R6928	T10659	T10647	meta	Spector,involves
R6929	T10773	T10769	cc	or,studies
R6930	T10774	T10775	compound	linkage,studies
R6931	T10775	T10769	conj	studies,studies
R6932	T10660	T10659	nmod	et,Spector
R6933	T10776	T10775	prep	in,studies
R6934	T10777	T10776	pobj	families,in
R6935	T10778	T10758	punct	.,linked
R6936	T10661	T10659	nmod	al.,Spector
R6937	T10780	T10781	nsubj	It,is
R6938	T10662	T10659	nummod	1996,Spector
R6939	T10782	T10781	acomp	interesting,is
R6940	T10663	T10659	punct	;,Spector
R6941	T10783	T10784	aux	to,note
R6942	T10664	T10659	nmod	Felson,Spector
R6943	T10784	T10781	xcomp	note,is
R6944	T10665	T10659	nmod	et,Spector
R6945	T10785	T10786	mark	that,contain
R6946	T10786	T10784	ccomp	contain,note
R6947	T10787	T10786	nsubj	several,contain
R6948	T10788	T10787	prep	of,several
R6949	T10666	T10659	nmod	al.,Spector
R6950	T10789	T10790	det	these,regions
R6951	T10790	T10788	pobj	regions,of
R6952	T10791	T10790	compound	chromosome,regions
R6953	T10667	T10659	nummod	1998,Spector
R6954	T10792	T10786	dobj	genes,contain
R6955	T10793	T10792	acl	encoding,genes
R6956	T10668	T10659	punct	;,Spector
R6957	T10794	T10795	amod	different,members
R6958	T10795	T10793	dobj	members,encoding
R6959	T10669	T10659	nmod	Hirsch,Spector
R6960	T10796	T10795	prep	of,members
R6961	T10797	T10798	det	the,pathway
R6962	T10798	T10796	pobj	pathway,of
R6963	T10670	T10659	nmod	et,Spector
R6964	T10799	T10800	compound	BMP,signaling
R6965	T10800	T10798	compound	signaling,pathway
R6966	T10801	T10792	punct	", ",genes
R6967	T10802	T10792	prep	including,genes
R6968	T10803	T10804	det	the,gene
R6969	T10804	T10802	pobj	gene,including
R6970	T10671	T10659	nmod	al.,Spector
R6971	T10805	T10804	compound	BMP5,gene
R6972	T10806	T10804	prep	on,gene
R6973	T10807	T10808	amod	human,6p12
R6974	T10672	T10659	nummod	1998,Spector
R6975	T10808	T10806	pobj	6p12,on
R6976	T10809	T10808	compound	chromosome,6p12
R6977	T10810	T10811	punct	(,Loughlin
R6978	T10673	T10659	punct	),Spector
R6979	T10811	T10804	meta	Loughlin,gene
R6980	T10812	T10811	nmod	et,Loughlin
R6981	T10674	T10647	punct	.,involves
R6982	T10813	T10811	nmod	al.,Loughlin
R6983	T10814	T10811	nummod	2002,Loughlin
R6984	T10815	T10811	punct	),Loughlin
R6985	T10676	T10677	nsubjpass	Humans,known
R6986	T10816	T10804	punct	", ",gene
R6987	T10817	T10818	det	the,gene
R6988	T10818	T10804	conj	gene,gene
R6989	T10819	T10818	compound	MADH1,gene
R6990	T10820	T10818	prep	on,gene
R6991	T10678	T10679	dep	that,are
R6992	T10821	T10822	amod	human,4q26
R6993	T10822	T10820	pobj	4q26,on
R6994	T10823	T10822	compound	chromosome,4q26
R6995	T10824	T10825	punct	–,4q31
R6996	T10679	T10676	relcl	are,Humans
R6997	T10825	T10822	prep	4q31,4q26
R6998	T10680	T10679	acomp	heterozygous,are
R6999	T10826	T10827	punct	(,Leppavuori
R7000	T10827	T10818	meta	Leppavuori,gene
R7001	T10828	T10827	nmod	et,Leppavuori
R7002	T10829	T10827	nmod	al.,Leppavuori
R7003	T10681	T10680	prep	for,heterozygous
R7004	T10830	T10827	nummod	1999,Leppavuori
R7005	T10831	T10827	punct	;,Leppavuori
R7006	T10682	T10683	nmod	loss,mutations
R7007	T10832	T10827	nmod	Kent,Leppavuori
R7008	T10833	T10827	nmod	et,Leppavuori
R7009	T10834	T10827	nmod	al.,Leppavuori
R7010	T10683	T10681	pobj	mutations,for
R7011	T10835	T10827	nummod	2002,Leppavuori
R7012	T10836	T10827	punct	),Leppavuori
R7013	T10837	T10818	punct	", ",gene
R7014	T10684	T10682	punct	-,loss
R7015	T10838	T10818	cc	and,gene
R7016	T10685	T10682	prep	of,loss
R7017	T10686	T10685	punct	-,of
R7018	T10687	T10685	pobj	function,of
R7019	T10839	T10840	det	the,receptor
R7020	T10688	T10683	prep	in,mutations
R7021	T10840	T10818	conj	receptor,gene
R7022	T10841	T10840	compound	BMPR2,receptor
R7023	T10842	T10840	prep	on,receptor
R7024	T10689	T10688	pobj	BMPR1A,in
R7025	T10843	T10844	amod	human,2q33
R7026	T10844	T10842	pobj	2q33,on
R7027	T10845	T10844	compound	chromosome,2q33
R7028	T10690	T10677	auxpass	are,known
R7029	T10846	T10847	punct	(,Wright
R7030	T10847	T10840	meta	Wright,receptor
R7031	T10848	T10847	nmod	et,Wright
R7032	T10691	T10692	aux	to,be
R7033	T10849	T10847	nmod	al.,Wright
R7034	T10850	T10847	nummod	1996,Wright
R7035	T10851	T10847	punct	),Wright
R7036	T10692	T10677	xcomp	be,known
R7037	T10852	T10781	punct	.,is
R7038	T10854	T10855	det	The,nature
R7039	T10693	T10692	prep	at,be
R7040	T10855	T10857	nsubj	nature,suggests
R7041	T10856	T10855	amod	complex,nature
R7042	T10694	T10693	pobj	risk,at
R7043	T10858	T10855	prep	of,nature
R7044	T10695	T10694	prep	for,risk
R7045	T10859	T10860	amod	human,osteoarthritis
R7046	T10860	T10858	pobj	osteoarthritis,of
R7047	T10861	T10862	mark	that,involved
R7048	T10696	T10697	amod	juvenile,polyposis
R7049	T10697	T10695	pobj	polyposis,for
R7050	T10862	T10857	ccomp	involved,suggests
R7051	T10698	T10699	punct	(,Howe
R7052	T10863	T10862	nsubjpass	interactions,involved
R7053	T10864	T10863	prep	between,interactions
R7054	T10865	T10866	amod	multiple,genes
R7055	T10699	T10677	meta	Howe,known
R7056	T10866	T10864	pobj	genes,between
R7057	T10867	T10862	aux	may,involved
R7058	T10868	T10862	auxpass	be,involved
R7059	T10700	T10699	nmod	et,Howe
R7060	T10869	T10862	prep	in,involved
R7061	T10870	T10869	pcomp	modifying,in
R7062	T10701	T10699	nmod	al.,Howe
R7063	T10871	T10870	dobj	susceptibility,modifying
R7064	T10872	T10871	prep	to,susceptibility
R7065	T10873	T10874	det	the,disease
R7066	T10702	T10699	nummod	2001,Howe
R7067	T10874	T10872	pobj	disease,to
R7068	T10875	T10857	punct	.,suggests
R7069	T10877	T10878	det	The,inclusion
R7070	T10878	T10879	nsubj	inclusion,help
R7071	T10703	T10699	punct	;,Howe
R7072	T10880	T10878	prep	of,inclusion
R7073	T10881	T10882	amod	genetic,markers
R7074	T10704	T10699	nmod	Zhou,Howe
R7075	T10882	T10880	pobj	markers,of
R7076	T10883	T10882	prep	near,markers
R7077	T10884	T10885	compound	BMP,components
R7078	T10705	T10699	nmod	et,Howe
R7079	T10885	T10883	pobj	components,near
R7080	T10886	T10885	compound	signaling,components
R7081	T10887	T10879	aux	may,help
R7082	T10706	T10699	nmod	al.,Howe
R7083	T10888	T10879	xcomp	identify,help
R7084	T10889	T10890	amod	additional,loci
R7085	T10890	T10888	dobj	loci,identify
R7086	T10707	T10699	nummod	2001,Howe
R7087	T10891	T10892	compound	osteoarthritis,susceptibility
R7088	T10892	T10890	compound	susceptibility,loci
R7089	T10708	T10699	punct	),Howe
R7090	T10893	T10888	cc	and,identify
R7091	T10894	T10888	conj	facilitate,identify
R7092	T10895	T10896	det	the,search
R7093	T10709	T10677	punct	", ",known
R7094	T10896	T10894	dobj	search,facilitate
R7095	T10897	T10896	prep	for,search
R7096	T10898	T10899	amod	causative,mutations
R7097	T10710	T10677	cc	but,known
R7098	T10899	T10897	pobj	mutations,for
R7099	T10900	T10879	punct	.,help
R7100	T10711	T10712	det	the,risk
R7101	T10902	T10903	nmod	Development,processes
R7102	T10903	T10906	nsubj	processes,been
R7103	T10712	T10713	nsubjpass	risk,reported
R7104	T10904	T10902	cc	and,Development
R7105	T10905	T10902	conj	disease,Development
R7106	T10713	T10677	conj	reported,known
R7107	T10907	T10903	prep	in,processes
R7108	T10908	T10909	amod	synovial,joints
R7109	T10714	T10712	prep	of,risk
R7110	T10909	T10907	pobj	joints,in
R7111	T10910	T10906	aux	have,been
R7112	T10911	T10906	acomp	difficult,been
R7113	T10715	T10714	pobj	osteoarthritis,of
R7114	T10912	T10913	aux	to,study
R7115	T10913	T10911	advcl	study,difficult
R7116	T10716	T10712	prep	for,risk
R7117	T10914	T10913	advmod	genetically,study
R7118	T10915	T10906	punct	", ",been
R7119	T10916	T10917	mark	because,generated
R7120	T10917	T10906	advcl	generated,been
R7121	T10918	T10919	amod	synovial,joints
R7122	T10919	T10917	nsubjpass	joints,generated
R7123	T10920	T10917	auxpass	are,generated
R7124	T10717	T10718	det	these,people
R7125	T10921	T10917	cc	and,generated
R7126	T10922	T10917	conj	function,generated
R7127	T10923	T10917	prep	at,generated
R7128	T10718	T10716	pobj	people,for
R7129	T10924	T10925	advmod	relatively,late
R7130	T10925	T10926	amod	late,stages
R7131	T10926	T10923	pobj	stages,at
R7132	T10719	T10713	aux	has,reported
R7133	T10927	T10926	prep	of,stages
R7134	T10928	T10929	compound	vertebrate,development
R7135	T10929	T10927	pobj	development,of
R7136	T10720	T10713	neg	not,reported
R7137	T10930	T10906	punct	.,been
R7138	T10721	T10713	auxpass	been,reported
R7139	T10932	T10933	det	The,system
R7140	T10933	T10937	nsubj	system,provides
R7141	T10934	T10935	compound	Gdf5,Cre
R7142	T10722	T10713	punct	.,reported
R7143	T10935	T10933	compound	Cre,system
R7144	T10936	T10935	punct	-,Cre
R7145	T10724	T10725	advmod	However,were
R7146	T10938	T10939	det	a,method
R7147	T10939	T10937	dobj	method,provides
R7148	T10940	T10939	amod	new,method
R7149	T10941	T10939	prep	for,method
R7150	T10726	T10725	punct	", ",were
R7151	T10942	T10941	pcomp	restricting,for
R7152	T10943	T10944	compound	gene,expression
R7153	T10944	T10942	dobj	expression,restricting
R7154	T10727	T10728	det	the,mice
R7155	T10728	T10725	nsubj	mice,were
R7156	T10729	T10728	compound	control,mice
R7157	T10945	T10944	cc	or,expression
R7158	T10730	T10728	acl	used,mice
R7159	T10946	T10944	conj	inactivation,expression
R7160	T10947	T10948	advmod	primarily,to
R7161	T10948	T10942	prep	to,restricting
R7162	T10731	T10730	prep	in,used
R7163	T10949	T10950	amod	articular,regions
R7164	T10950	T10948	pobj	regions,to
R7165	T10732	T10733	det	this,study
R7166	T10951	T10942	punct	", ",restricting
R7167	T10952	T10953	advmod	thus,avoiding
R7168	T10953	T10942	advcl	avoiding,restricting
R7169	T10954	T10955	det	the,functions
R7170	T10955	T10953	dobj	functions,avoiding
R7171	T10956	T10955	amod	pleiotropic,functions
R7172	T10957	T10955	prep	of,functions
R7173	T10958	T10959	amod	many,genes
R7174	T10959	T10957	pobj	genes,of
R7175	T10960	T10955	prep	in,functions
R7176	T10961	T10962	amod	other,tissues
R7177	T10962	T10960	pobj	tissues,in
R7178	T10963	T10937	punct	.,provides
R7179	T10965	T10966	prep	Depending,used
R7180	T10967	T10965	prep	on,Depending
R7181	T10968	T10969	det	the,configuration
R7182	T10969	T10967	pobj	configuration,on
R7183	T10970	T10969	prep	of,configuration
R7184	T10971	T10972	det	the,gene
R7185	T10972	T10970	pobj	gene,of
R7186	T10973	T10972	amod	floxed,gene
R7187	T10974	T10972	compound	target,gene
R7188	T10975	T10966	punct	", ",used
R7189	T10976	T10977	det	this,system
R7190	T10977	T10966	nsubjpass	system,used
R7191	T10978	T10966	aux	can,used
R7192	T10979	T10966	auxpass	be,used
R7193	T10980	T10981	aux	to,activate
R7194	T10981	T10966	advcl	activate,used
R7195	T10982	T10981	preconj	either,activate
R7196	T10983	T10984	det	the,expression
R7197	T10984	T10981	dobj	expression,activate
R7198	T10985	T10984	prep	of,expression
R7199	T10986	T10987	det	a,gene
R7200	T10987	T10985	pobj	gene,of
R7201	T10988	T10989	advmod	primarily,in
R7202	T10989	T10981	prep	in,activate
R7203	T10990	T10991	amod	developing,joints
R7204	T10991	T10989	pobj	joints,in
R7205	T10992	T10993	punct	(,ssee
R7206	T10993	T10981	parataxis	ssee,activate
R7207	T10994	T10995	nmod	Figure,1B
R7208	T10995	T10993	dobj	1B,ssee
R7209	T10996	T10997	punct	–,1D
R7210	T10997	T10995	prep	1D,1B
R7211	T10998	T10993	punct	),ssee
R7212	T10999	T10981	punct	", ",activate
R7213	T11000	T10981	cc	or,activate
R7214	T11001	T11002	aux	to,inactivate
R7215	T11002	T10981	conj	inactivate,activate
R7216	T11003	T11004	compound	gene,function
R7217	T11004	T11002	dobj	function,inactivate
R7218	T11005	T11002	prep	in,inactivate
R7219	T11006	T11007	amod	articular,regions
R7220	T11007	T11005	pobj	regions,in
R7221	T11008	T11009	punct	(,see
R7222	T11009	T10966	parataxis	see,used
R7223	T11010	T11009	dobj	Figure,see
R7224	T11011	T11010	nummod	3,Figure
R7225	T11012	T11009	punct	),see
R7226	T11013	T10966	punct	.,used
R7227	T11015	T11016	amod	Additional,studies
R7228	T11016	T11017	nsubj	studies,enhance
R7229	T11018	T11016	prep	with,studies
R7230	T11019	T11020	det	this,system
R7231	T11020	T11018	pobj	system,with
R7232	T11021	T11017	aux	should,enhance
R7233	T11022	T11017	advmod	greatly,enhance
R7234	T11023	T11024	poss	our,knowledge
R7235	T11024	T11017	dobj	knowledge,enhance
R7236	T11025	T11024	prep	of,knowledge
R7237	T11026	T11027	det	the,mechanisms
R7238	T11027	T11025	pobj	mechanisms,of
R7239	T11028	T11027	nmod	development,mechanisms
R7240	T11029	T11028	punct	", ",development
R7241	T11030	T11028	conj	function,development
R7242	T11031	T11030	punct	", ",function
R7243	T11032	T11030	cc	and,function
R7244	T11033	T11030	conj	disease,function
R7245	T11034	T11027	prep	of,mechanisms
R7246	T11035	T11034	pobj	joints,of
R7247	T11036	T11017	punct	", ",enhance
R7248	T11037	T11017	cc	and,enhance
R7249	T11038	T11039	aux	may,bring
R7250	T11039	T11017	conj	bring,enhance
R7251	T11040	T11039	dobj	us,bring
R7252	T11041	T11039	advcl	closer,bring
R7253	T11042	T11041	prep	to,closer
R7254	T11043	T11044	amod	better,prevention
R7255	T11044	T11042	pobj	prevention,to
R7256	T11045	T11044	cc	and,prevention
R7257	T11046	T11044	conj	treatment,prevention
R7258	T11047	T11044	prep	of,prevention
R7259	T11048	T11049	compound	joint,diseases
R7260	T11049	T11047	pobj	diseases,of
R7261	T11050	T11017	punct	.,enhance

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PMC:523229 / 40394-50514 JSON TXT 5 Projects