Nuclei are intricately structured, and nuclear metabolism has an elaborate spatial organization. The architecture of the nucleus includes two overlapping and nucleic-acid-containing structures - chromatin and a nuclear matrix. The nuclear matrix is observed by microscopy in live, fixed and extracted cells. Its ultrastructure and composition show it to be, in large part, the ribonucleoprotein (RNP) network first seen in unfractionated cells more than 30 years ago. At that time, the discovery of this RNP structure explained surprising observations that RNA, packaged in proteins, is attached to an intranuclear, non-chromatin structure. Periodic and specific attachments of chromatin fibers to the nuclear matrix create the chromatin loop domains that can be directly observed by microscopy or inferred from biochemical experiments. The ultrastructure of the nuclear matrix is well characterized and consists of a nuclear lamina and an internal nuclear network of subassemblies linked together by highly structured fibers. These complex fibers are built on an underlying scaffolding of branched 10-nm filaments that connect to the nuclear lamina. The structural proteins of the nuclear lamina have been well characterized, but the structural biochemistry of the internal nuclear matrix has received less attention. Many internal matrix proteins have been identified, but far less is known about how these proteins assemble to make the fibers, filaments and other assemblies of the internal nuclear matrix. Correcting this imbalance will require the combined application of biochemistry and electron microscopy. The central problem in trying to define nuclear matrix structure is to identify the proteins that assemble into the 10-nm filaments upon which the interior architecture of the nucleus is constructed. Only by achieving a biochemical characterization of the nuclear matrix will we advance beyond simple microscopic observations of structure to a better understanding of nuclear matrix function, regulation and post-mitotic assembly.
REFERENCES
Abney
J. R.
, Cutler
B.
, Fillbach
M. L.
, Axelrod
D.
, Scalettar
B. A.
(1997
). Chromatin dynamics in interphase nuclei and its implications for nuclear structure.
J. Cell Biol
137
, 1459
–1468
Adolph
K. W.
, Cheng
S. M.
, Paulson
J. R.
, Laemmli
U. K.
(1977
). Isolation of a protein scaffold from mitotic HeLa cell chromosomes.
Proc. Nat. Acad. Sci. USA
74
, 4937
–4941
Adolph
K. W.
(1980
). Organization of chromosomes in HeLa cells: isolation of histone-depleted nuclei and nuclear scaffolds.
J. Cell Sci
42
, 291
–304
Alvarez
J. D.
, Yasui
D. H.
, Niida
H.
, Joh
T.
, Loh
D. Y.
, Kohwi-Shigematsu
T.
(2000
). The MAR-binding protein SATB1 orchestrates temporal and spatial expression of multiple genes during T-cell development.
Genes Dev
14
, 521
–535
Amankwah
K. S.
, De Boni
U.
(1994
). Ultrastructural localization of filamentous actin within neuronal interphase nuclei in situ.
Exp. Cell Res
210
, 315
–325
Bachellerie
J. P.
, Puvion
E.
, Zalta
J. P.
(1975
). Ultrastructural organization and biochemical characterization of chromatin- RNA- protein complexes isolated from mammalian cell nuclei.
Eur. J. Biochem
58
, 327
–337
Benyajati
C.
, Worcel
A.
(1976
). Isolation, characterization, and structure of the folded interphase genome of Drosophila melanogaster.
Cell
9
, 393
–407
Berezney
R.
, Coffey
D. S.
(1974
). Identification of a nuclear protein matrix.
Biochem. Biophys. Res. Commun
60
, 1410
–1417
Berezney
R.
(1980
). Fractionation of the nuclear matrix. I. Partial separation into matrix protein fibrils and a residual ribonucleoprotein fraction.
J. Cell Biol
85
, 641
–650
Bernhard
W.
(1969
). A new staining procedure for electron microscopical cytology.
J. Ultrastruct. Res
27
, 250
–265
Biggiogera
M.
, Fakan
S.
(1998
). Fine structural specific visualization of RNA on ultrathin sections.
J. Histochem. Cytochem
46
, 389
–395
Blencowe
B. J.
, Bauren
G.
, Eldridge
A. G.
, Issner
R.
, Nickerson
J. A.
, Rosonina
E.
, Sharp
P. A.
(2000
). The SRm160/300 splicing coactivator subunits.
RNA
6
, 111
–120
Blencowe
B. J.
, Nickerson
J. A.
, Issner
R.
, Penman
S.
, Sharp
P. A.
(1994
). Association of nuclear matrix antigens with exon-containing splicing complexes.
J. Cell Biol
127
, 593
–607
Blencowe
B. J.
, Issner
R.
, Nickerson
J. A.
, Sharp
P. A.
(1998
). A coactivator of pre-mRNA splicing.
Genes Dev
12
, 996
–1009
Bode
J.
, Kohwi
Y.
, Dickinson
L.
, Joh
T.
, Klehr
D.
, Mielke
C.
, Kohwi-Shigematsu
T.
(1992
). Biological significance of unwinding capability of nuclear matrix-associating DNAs.
Science
255
, 195
–197
Bornfleth
H.
, Edelmann
P.
, Zink
D.
, Cremer
T.
, Cremer
C.
(1999
). Quantitative motion analysis of subchromosomal foci in living cells using four-dimensional microscopy.
Biophys. J
77
, 2871
–2886
Boulikas
T.
(1995
). Chromatin domains and prediction of MAR sequences.
Int. Rev. Cytol
162
, 279
–388
Brasch
K.
(1982
). Fine structure and localization of the nuclear matrix in situ.
Exp. Cell Res
140
, 161
–171
Capco
D. G.
, Wan
K. M.
, Penman
S.
(1982
). The nuclear matrix: three-dimensional architecture and protein composition.
Cell
29
, 847
–858
Clemson
C. M.
, McNeil
J. A.
, Willard
H. F.
, Lawrence
J. B.
(1996
). XIST RNA paints the inactive X chromosome at interphase: evidence for a novel RNA involved in nuclear/chromosome structure.
J. Cell Biol
132
, 259
–275
Cmarko
D.
, Verschure
P. J.
, Martin
T. E.
, Dahmus
M. E.
, Krause
S.
, Fu
X. D.
, van Driel
R.
, Fakan
S.
(1999
). Ultrastructural analysis of transcription and splicing in the cell nucleus after bromo-UTP microinjection.
Mol. Biol. Cell
10
, 211
–223
Cook
P. R.
, Brazell
I. A.
, Jost
E.
(1976
). Characterization of nuclear structures containing superhelical DNA.
J. Cell Sci
22
, 303
–324
Cook
P. R.
(1999
). The organization of replication and transcription.
Science
284
, 1790
–1795
Cremer
C.
, Munkel
C.
, Granzow
M.
, Jauch
A.
, Dietzel
S.
, Eils
R.
, Guan
X. Y.
, Meltzer
P. S.
, Trent
J. M.
, Langowski
J.
, et al. (1996
). Nuclear architecture and the induction of chromosomal aberrations.
Mutat. Res
366
, 97
–116
de Belle
I.
, Cai
S.
, Kohwi-Shigematsu
T.
(1998
). The genomic sequences bound to special AT-rich sequence-binding protein 1 (SATB1) in vivo in Jurkat T cells are tightly associated with the nuclear matrix at the bases of the chromatin loops.
J. Cell Biol
141
, 335
–348
Derenzini
M.
, Marinozzi
V.
, Novello
F.
(1976
). Effects of alpha-amanitine on chromatin in regenerating rat hepatocytes: a biochemical and morphologic study.
Virchows Arch. B Cell. Pathol
20
, 307
–318
Derenzini
M.
, Pession-Brizzi
A.
, Betts-Eusebi
C.
, Novello
F.
(1981
). Relationship between the fine structural organization of chromatin and nucleic acid synthesis in regenerating rat hepatocytes.
J. Ultrastruct. Res
75
, 229
–242
Dickinson
P.
, Cook
P. R.
, Jackson
D. A.
(1990
). Active RNA polymerase I is fixed within the nucleus of HeLa cells.
EMBO J
9
, 2207
–2214
Dickinson
L. A.
, Joh
T.
, Kohwi
Y.
, Kohwi-Shigematsu
T.
(1992
). A tissue-specific MAR/SAR DNA-binding protein with unusual binding site recognition.
Cell
70
, 631
–645
Engelhardt
M.
(1999
). Demonstration of a DNase-sensitive network associated with the nuclear pore complexes in rat liver nuclei.
Chromosoma
108
, 64
–71
Faiferman
I.
, Pogo
A. O.
(1975
). Isolation of a nuclear ribonucleoprotein network that contains heterogeneous RNA and is bound to the nuclear envelope.
Biochemistry
14
, 3808
–3816
Fey
E. G.
, Ornelles
D. A.
, Penman
S.
(1986
). Association of RNA with the cytoskeleton and the nuclear matrix.
J. Cell Sci. Suppl
5
, 99
–119
Fey
E. G.
, Penman
S.
(1988
). Nuclear matrix proteins reflect cell typeof origin in cultured human cells.
Proc. Nat. Acad. Sci. USA
85
, 121
–125
Gallinaro
H.
, Puvion
E.
, Kister
L.
, Jacob
M.
(1983
). Nuclear matrixand hnRNP share a common structural constituent associated with premessenger RNA.
EMBO J
2
, 953
–960
Gerdes
M. G.
, Carter
K. C.
, Moen
P. T.
Jr., Lawrence
J. B.
(1994
). Dynamic changes in the higher-level chromatin organization of specific sequences revealed by in situ hybridization to nuclear halos.
J. Cell Biol
126
, 289
–304
Getzenberg
R. H.
, Pienta
K. J.
, Huang
E. Y.
, Coffey
D. S.
(1991
). Identification of nuclear matrix proteins in the cancer and normal rat prostate.
Cancer Res
51
, 6514
–6520
Goldberg
M.
, Harel
A.
, Gruenbaum
Y.
(1999
). The nuclear lamina: molecular organization and interaction with chromatin.
Crit. Rev. Eukaryot. Gene Expr
9
, 285
–293
Gruenbaum
Y.
, Wilson
K. L.
, Harel
A.
, Goldberg
M.
, Cohen
M.
(2000
). Review: nuclear lamins—structural proteins with fundamental functions.
J. Struct. Biol
129
, 313
–323
Gueth-Hallonet
C.
, Wang
J.
, Harborth
J.
, Weber
K.
, Osborn
M.
(1998
). Induction of a regular nuclear lattice by overexpression of NuMA.
Exp. Cell Res
243
, 434
–452
Harborth
J.
, Wang
J.
, Gueth-Hallonet
C.
, Weber
K.
, Osborn
M.
(1999
). Self assembly of NuMA: multiarm oligomers as structural units of a nuclear lattice.
EMBO J
18
, 1689
–1700
He
D. C.
, Nickerson
J. A.
, Penman
S.
(1990
). Core filaments of the nuclear matrix.
J. Cell Biol
110
, 569
–580
He
D. C.
, Martin
T.
, Penman
S.
(1991
). Localization of heterogeneous nuclear ribonucleoprotein in the interphase nuclear matrix core filaments and on perichromosomal filaments at mitosis.
Proc. Nat. Acad. Sci. USA
88
, 7469
–7473
Hendzel
M. J.
, Sun
J. M.
, Chen
H. Y.
, Rattner
J. B.
, Davie
J. R.
(1994
). Histone acetyltransferase is associated with the nuclear matrix.
J. Biol. Chem
269
, 22894
–22901
Hendzel
M. J.
, Boisvert
F.
, Bazett-Jones
D. P.
(1999
). Direct visualization of a protein nuclear architecture.
Mol. Biol. Cell
10
, 2051
–2062
Herman
R.
, Weymouth
L.
, Penman
S.
(1978
). Heterogeneous nuclear RNA-protein fibers in chromatin-depleted nuclei.
J. Cell Biol
78
, 663
–674
Hozak
P.
, Hassan
A. B.
, Jackson
D. A.
, Cook
P. R.
(1993
). Visualization of replication factories attached to nucleoskeleton.
Cell
73
, 361
–373
Hozak
P.
, Sasseville
A. M.
, Raymond
Y.
, Cook
P. R.
(1995
). Lamin proteins form an internal nucleoskeleton as well as a peripheral lamina in human cells.
J. Cell Sci
108
, 635
–644
Jackson
D. A.
, Cook
P. R.
(1985
). Transcription occurs at a nucleoskeleton.
EMBO J
4
, 919
–925
Jackson
D. A.
, Cook
P. R.
(1988
). Visualization of a filamentous nucleoskeleton with a 23 nm axial repeat.
EMBO J
7
, 3667
–3677
Jackson
D. A.
, Dickinson
P.
, Cook
P. R.
(1990
). The size of chromatin loops in HeLa cells.
EMBO J
9
, 567
–571
Jackson
D. A.
, Iborra
F. J.
, Manders
E. M.
, Cook
P. R.
(1998
). Numbers and organization of RNA polymerases, nascent transcripts, and transcription units in HeLa nuclei.
Mol. Biol. Cell
9
, 1523
–1536
Johansen
K. M.
, Johansen
J.
, Jin
Y.
, Walker
D. L.
, Wang
D.
, Wang
Y.
(1999
). Chromatin structure and nuclear remodeling.
Crit. Rev. Euk. Gene Exp
9
, 267
–277
Kaufmann
S. H.
, Shaper
J. H.
(1984
). A subset of non-histone nuclear proteins reversibly stabilized by the sulfhydryl cross-linking reagent tetrathionate. Polypeptides of the internal nuclear matrix.
Exp. Cell Res
155
, 477
–495
Krauss
S. W.
, Larabell
C. A.
, Lockett
S.
, Gascard
P.
, Penman
S.
, Mohandas
N.
, Chasis
J. A.
(1997
). Structural protein 4. 1 in the nucleus of human cells: dynamic rearrangements during cell division.
J. Cell Biol
137
, 275
–289
Kruhlak
M. J.
, Lever
M. A.
, Fischle
W.
, Verdin
E.
, Bazett-Jones
D. P.
, Hendzel
M. J.
(2000
). Reduced Mobility of the Alternate Splicing Factor (ASF) through the Nucleoplasm and Steady State Speckle Compartments.
J. Cell Biol
150
, 41
–52
Kurz
A.
, Lampel
S.
, Nickolenko
J. E.
, Bradl
J.
, Benner
A.
, Zirbel
R.M.
, Cremer
T.
, Lichter
P.
(1996
). Active and inactive genes localize preferentially in the periphery of chromosome territories.
J. Cell Biol
135
, 1195
–205
Leonhardt
H.
, Rahn
H. P.
, Weinzierl
P.
, Sporbert
A.
, Cremer
T.
, Zink
D.
, Cardoso
M. C.
(2000
). Dynamics of DNA replication factories in living cells.
J. Cell Biol
149
, 271
–280
Leser
G. P.
, Escara-Wilke
J.
, Martin
T. E.
(1984
). Monoclonal antibodies to heterogeneous nuclear RNA-protein complexes. The core proteins comprise a conserved group of related polypeptides.
J. Biol. Chem
259
, 1827
–1833
Ma
H.
, Siegel
A. J.
, Berezney
R.
(1999
). Association of chromosome territories with the nuclear matrix. Disruption of human chromosome territories correlates with the release of a subset of nuclear matrix proteins.
J. Cell Biol
146
, 531
–542
Mancini
M. G.
, Liu
B.
, Sharp
Z. D.
, Mancini
M. A.
(1999
). Subnuclear partitioning and functional regulation of the Pit-1 transcription factor.
J. Cell. Biochem
72
, 322
–338
Maniotis
A. J.
, Bojanowski
K.
, Ingber
D. E.
(1997
). Mechanical continuity and reversible chromosome disassembly within intact genomes removed from living cells.
J. Cell. Biochem
65
, 114
–130
Mattagajasingh
S. N.
, Huang
S. C.
, Hartenstein
J. S.
, Snyder
M.
, Marchesi
V. T.
, Benz
E. J.
(1999
). A nonerythroid isoform of protein 4. 1R interacts with the nuclear mitotic apparatus (NuMA) protein.
J. Cell Biol
145
, 29
–43
Mattern
K. A.
, Humbel
B. M.
, Muijsers
A. O.
, de Jong
L.
, van Driel
R.
(1996
). hnRNP proteins and B23 are the major proteins of the internal nuclear matrix of HeLa S3 cells.
J. Cell. Biochem
62
, 275
–289
McNally
J. G.
, Muller
W. G.
, Walker
D.
, Wolford
R.
, Hager
G. L.
(2000
). The glucocorticoid receptor: rapid exchange with regulatory sites in living cells.
Science
287
, 1262
–1265
Mintz
P. J.
, Patterson
S. D.
, Neuwald
A. F.
, Spahr
C. S.
, Spector
D. L.
(1999
). Purification and biochemical characterization of interchromatin granule clusters.
EMBO J
18
, 4308
–4320
Miralles
F.
, Ofverstedt
L. G.
, Sabri
N.
, Aissouni
Y.
, Hellman
U.
, Skoglund
U.
, Visa
N.
(2000
). Electron tomography reveals posttranscriptional binding of pre-mRNPs to specific fibers in the nucleoplasm.
J. Cell Biol
148
, 271
–282
Mirkovitch
J.
, Mirault
M. E.
, Laemmli
U. K.
(1984
). Organization of the higher-order chromatin loop: specific DNA attachment sites on nuclear scaffold.
Cell
39
, 223
–232
Mirkovitch
J.
, Spierer
P.
, Laemmli
U. K.
(1986
). Genes and loops in 320, 000 base-pairs of the Drosophila melanogaster chromosome.
J. Mol. Biol
190
, 255
–258
Misteli
T.
, Spector
D. L.
(1996
). Serine/threonine phosphatase 1 modulates the subnuclear distribution of pre-mRNA splicing factors.
Mol. Biol. Cell
7
, 1559
–1572
Misteli
T.
, Caceres
J. F.
, Spector
D. L.
(1997
). The dynamics of a pre-mRNA splicing factor in living cells.
Nature
387
, 523
–527
Misteli
T.
, Spector
D. L.
(1998
). The cellular organization of gene expression.
Curr. Opin. Cell Biol
10
, 323
–331
Misteli
T.
(2000
). Cell biology of transcription and pre-mRNA splicing: nuclear architecture meets nuclear function.
J. Cell Sci
113
, 1841
–1849
Monneron
A.
, Bernhard
W.
(1969
). Fine structural organization of the interphase nucleus in some mammalian cells.
J. Ultrastruct. Res
27
, 266
–288
Nakayasu
H.
, Ueda
K.
(1985
). Association of rapidly-labelled RNAs with actin in nuclear matrix from mouse L5178Y cells.
Exp. Cell Res
160
, 319
–330
Nakayasu
H.
, Berezney
R.
(1989
). Mapping replicational sites in the eucaryotic cell nucleus.
J. Cell Biol
108
, 1
–11
Narayan
K. S.
, Steele
W. J.
, Smetana
K.
, Busch
H.
(1967
). Ultrastructural aspects of the ribonucleo-protein network in nuclei of Walker tumor and rat liver.
Exp. Cell Res
46
, 65
–77
Nelson
W. G.
, Pienta
K. J.
, Barrack
E. R.
, Coffey
D. S.
(1986
). The role of the nuclear matrix in the organization and function of DNA.
Annu. Rev. Biophys. Biophys. Chem
15
, 457
–475
Nickerson
J. A.
, Krochmalnic
G.
, Wan
K. M.
, Penman
S.
(1989
). Chromatin architecture and nuclear RNA.
Proc. Nat. Acad. Sci. USA
86
, 177
–181
Nickerson
J. A.
, Krockmalnic
G.
, Wan
K. M.
, Penman
S.
(1997
). The nuclear matrix revealed by eluting chromatin from a cross-linked nucleus.
Proc. Nat. Acad. Sci. USA
94
, 4446
–4450
Nowak
G.
, Pestic-Dragovich
L.
, Hozak
P.
, Philimonenko
A.
, Simerly
C.
, Schatten
G.
, de Lanerolle
P.
(1997
). Evidence for the presence of myosin I in the nucleus.
J. Biol. Chem
272
, 17176
–17181
Oegema
K.
, Marshall
W. F.
, Sedat
J. W.
, Alberts
B. M.
(1997
). Two proteins that cycle asynchronously between centrosomes and nuclear structures: Drosophila CP60 and CP190.
J. Cell Sci
110
, 1573
–1583
Panning
B.
, Jaenisch
R.
(1998
). RNA and the epigenetic regulation of X chromosome inactivation.
Cell
93
, 305
–308
Pardoll
D. M.
, Vogelstein
B.
, Coffey
D. S.
(1980
). A fixed site of DNA replication in eucaryotic cells.
Cell
19
, 527
–536
Pederson
T.
(2000
). Half a century of ‘the nuclear matrix’.
Mol. Biol. Cell
11
, 799
–805
Phair
R. D.
, Misteli
T.
(2000
). High mobility of proteins in the mammalian cell nucleus.
Nature
404
, 604
–609
Politz
J. C.
, Browne
E. S.
, Wolf
D. E.
, Pederson
T.
(1998
). Intranuclear diffusion and hybridization state of oligonucleotides measured by fluorescence correlation spectroscopy in living cells.
Proc. Nat. Acad. Sci. USA
95
, 6043
–6048
Politz
J. C.
, Tuft
R. A.
, Pederson
T.
, Singer
R. H.
(1999
). Movement of nuclear poly(A) RNA throughout the interchromatin space in living cells.
Curr. Biol
9
, 285
–291
Reyes
J. C.
, Muchardt
C.
, Yaniv
M.
(1997
). Components of the human SWI/SNF complex are enriched in active chromatin and are associated with the nuclear matrix.
J. Cell Biol
137
, 263
–274
Samarina
O. P.
, Lukanidin
E. M.
, Molnar
J.
, Georgiev
G. P.
(1968
). Structural organization of nuclear complexes containing DNA-like RNA.
J. Mol. Biol
33
, 251
–263
Schroder
H. C.
, Trolltsch
D.
, Wenger
R.
, Bachmann
M.
, Diehl-Seifert
B.
, Muller
W. E.
(1987
). Cytochalasin B selectively releases ovalbumin mRNA precursors but not the mature ovalbumin mRNA from hen oviduct nuclear matrix.
Eur. J. Biochem
167
, 239
–245
Shopland
L. S.
, Lawrence
J. B.
(2000
). Seeking Common Ground in Nuclear Complexity.
J. Cell Biol
150
, 1
–.Singh
O. P.
, Bjorkroth
B.
, Masich
S.
, Wieslander
L.
, Daneholt
B.
(1999
). The intranuclear movement of Balbiani ring premessenger ribonucleoprotein particles.
Exp. Cell Res
251
, 135
–146
Smetana
K.
, Steele
W. J.
, Busch
H.
(1963
). A nuclear ribonucleoprotein network.
Exp. Cell Res
31
, 198
–201
Smith
H. C.
, Puvion
E.
, Buchholtz
L. A.
, Berezney
R.
(1984
). Spatial distribution of DNA loop attachment and replicational sites in the nuclear matrix.
J. Cell Biol
99
, 1794
–1802
Smith
K. P.
, Moen
P. T.
, Wydner
K. L.
, Coleman
J. R.
, Lawrence
J. B.
(1999
). Processing of endogenous pre-mRNAs in association with SC-35 domains is gene specific.
J. Cell Biol
144
, 617
–629
Spector
D. L.
(1996
). Nuclear organization and gene expression.
Exp. Cell Res
229
, 189
–197
Stenoien
D. L.
, Mancini
M. G.
, Patel
K.
, Allegretto
E. A.
, Smith
C. L.
, Mancini
M. A.
(2000
). Subnuclear trafficking of estrogen receptor-alpha and steroid receptor coactivator-1.
Mol. Endocrinol
14
, 518
–534
Sun
J. M.
, Chen
H. Y.
, Moniwa
M.
, Samuel
S.
, Davie
J. R.
(1999
). Purification and characterization of chicken erythrocyte histone deacetylase 1.
Biochemistry
38
, 5939
–5947
Tan
J.
, Wooley
J. C.
, LeStourgeon
W. M.
(2000
). Nuclear matrix-like filaments and fibrogranular complexes form through the rearrangement of specific nuclear ribonucleoproteins.
Mol. Biol. Cell
11
, 1547
–1554
Tubo
R. A.
, Berezney
R.
(1987
). Nuclear matrix-bound DNA primase. Elucidation of an RNA priming system in nuclear matrix isolated from regenerating rat liver.
J. Biol. Chem
262
, 6637
–6642
Tubo
R. A.
, Martelli
A. M.
, Berezney
R.
(1987
). Enhanced processivity of nuclear matrix bound DNA polymerase alpha from regenerating rat liver.
Biochemistry
26
, 5710
–5718
van Venrooij
W. J.
, Janssen
D. B.
(1978
). HnRNP particles.
Mol. Biol. Rep
4
, 3
–8
van Wijnen
A. J.
, Bidwell
J. P.
, Fey
E. G.
, Penman
S.
, Lian
J. B.
, Stein
J. L.
, Stein
G. S.
(1993
). Nuclear matrix association of multiple sequence-specific DNA binding activities related to SP-1, ATF, CCAAT, C/EBP, OCT-1, and AP-1.
Biochemistry
32
, 8397
–8402
Vogelstein
B.
, Pardoll
D. M.
, Coffey
D. S.
(1980
). Supercoiled loops and eucaryotic DNA replicaton.
Cell
22
, 79
–85
Walker
B. W.
, Lothstein
L.
, Baker
C. L.
, LeStourgeon
W. M.
(1980
). The release of 40S hnRNP particles by brief digestion of HeLa nuclei with micrococcal nuclease.
Nucl. Acids Res
8
, 3639
–3657
Wan
K. M.
, Nickerson
J. A.
, Krockmalnic
G.
, Penman
S.
(1999
). The nuclear matrix prepared by amine modification.
Proc. Nat. Acad. Sci. USA
96
, 933
–938
Wansink
D. G.
, Schul
W.
, van der Kraan
I.
, van Steensel
B.
, van Driel
R.
, de Jong
L.
(1993
). Fluorescent labeling of nascent RNA reveals transcription by RNA polymerase II in domains scattered throughout the nucleus.
J. Cell Biol
122
, 283
–293
Wansink
D. G.
, Sibon
O. C.
, Cremers
F. F.
, van Driel
R.
, de Jong
L.
(1996
). Ultrastructural localization of active genes in nuclei of A431 cells.
J. Cell. Biochem
62
, 10
–18
Wei
X.
, Somanathan
S.
, Samarabandu
J.
, Berezney
R.
(1999
). Three-dimensional visualization of transcription sites and their associationwith splicing factor-rich nuclear speckles.
J. Cell Biol
146
, 543
–558
Xing
Y. G.
, Lawrence
J. B.
(1991
). Preservation of specific RNA distribution within the chromatin-depleted nuclear substructure demonstrated by in situ hybridization coupled with biochemical fractionation.
J. Cell Biol
112
, 1055
–1063
Zeitlin
S.
, Parent
A.
, Silverstein
S.
, Efstratiadis
A.
(1987
). Pre-mRNA splicing and the nuclear matrix.
Mol. Cell Biol
7
, 111
–120
Zeitlin
S.
, Wilson
R. C.
, Efstratiadis
A.
(1989
). Autonomous splicing and complementation of in vivo-assembled spliceosomes.
J. Cell Biol
108
, 765
–777
Zeng
C.
, van Wijnen
A. J.
, Stein
J. L.
, Meyers
S.
, Sun
W.
, Shopland
L.
, Lawrence
J. B.
, Penman
S.
, Lian
J. B.
, Stein
G. S.
, et al. (1997
). Identification of a nuclear matrix targeting signal in the leukemia and bone-related AML/CBF-alpha transcription factors.
Proc. Nat. Acad. Sci. USA
94
, 6746
–6751
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