When is dna condensed during the cell cycle




















Cell 21, 51— Dultz, E. Systematic kinetic analysis of mitotic dis- and reassembly of the nuclear pore in living cells. Egli, D. Mediators of reprogramming: transcription factors and transitions through mitosis. Farrell, J. Embryonic onset of late replication requires Cdc25 down-regulation.

Ferreira, H. Histone modifications influence the action of Snf2 family remodelling enzymes by different mechanisms. Filippakopoulos, P. Targeting bromodomains: epigenetic readers of lysine acetylation. Drug Discov. Histone recognition and large-scale structural analysis of the human bromodomain family. Finlan, L. Recruitment to the nuclear periphery can alter expression of genes in human cells. PLoS Genet. Fischle, W. Regulation of HP1-chromatin binding by histone H3 methylation and phosphorylation.

Francis, N. Fuchs, E. The tortoise and the hair: slow-cycling cells in the stem cell race. A matter of life and death: self-renewal in stem cells. EMBO Rep. Funakoshi, T. Localization of Pom to the inner nuclear membrane is required for an early step of interphase nuclear pore complex assembly.

Cell 22, — Ganier, O. Synergic reprogramming of mammalian cells by combined exposure to mitotic Xenopus egg extracts and transcription factors. Gaydos, L. Gene repression. H3K27me and PRC2 transmit a memory of repression across generations and during development. Gialitakis, M.

Gamma interferon-dependent transcriptional memory via relocalization of a gene locus to PML nuclear bodies. Giet, R. Drosophila aurora B kinase is required for histone H3 phosphorylation and condensin recruitment during chromosome condensation and to organize the central spindle during cytokinesis.

Goren, A. DNA replication timing of the human beta-globin domain is controlled by histone modification at the origin. Gottesfeld, J. Mitotic repression of the transcriptional machinery. Trends Biochem. Groth, A. Regulation of replication fork progression through histone supply and demand.

Chromatin challenges during DNA replication and repair. Guelen, L. Domain organization of human chromosomes revealed by mapping of nuclear lamina interactions. Gunesdogan, U. Histone supply regulates S phase timing and cell cycle progression. Elife 3:e Guttinger, S. Orchestrating nuclear envelope disassembly and reassembly during mitosis. Halley-Stott, R. Mitosis gives a brief window of opportunity for a change in gene transcription.

PLoS Biol. Hanna, J. Direct cell reprogramming is a stochastic process amenable to acceleration. Hansen, K. A model for transmission of the H3K27me3 epigenetic mark.

Hansen, R. Sequencing newly replicated DNA reveals widespread plasticity in human replication timing. Hardwick, L. Nervous decision-making: to divide or differentiate. Hargreaves, D. ATP-dependent chromatin remodeling: genetics, genomics and mechanisms. Cell Res. Hayashi, M. Henley, S. The retinoblastoma family of proteins and their regulatory functions in the mammalian cell division cycle. Cell Div. Hentschel, C. The organization and expression of histone gene families. Hetzer, M.

Border control at the nucleus: biogenesis and organization of the nuclear membrane and pore complexes. Cell 17, — Hirano, T. Condensins: universal organizers of chromosomes with diverse functions. Ho, J. Comparative analysis of metazoan chromatin organization. Houston, S. Catalytic function of the PR-Set7 histone H4 lysine 20 monomethyltransferase is essential for mitotic entry and genomic stability.

Hsiung, C. Genome accessibility is widely preserved and locally modulated during mitosis. Iizuka, M. Regulation of replication licensing by acetyltransferase Hbo1. Imamoto, N. Nuclear pore dynamics during the cell cycle. Ito, S. Jenuwein, T. Translating the histone code. Jin, S. Genomic mapping of 5-hydroxymethylcytosine in the human brain. Nucleic Acids Res. Joo, H. Regulation of cell cycle progression and gene expression by H2A deubiquitination.

Kadauke, S. Mitotic bookmarking by transcription factors. Epigenetics Chromatin Tissue-specific mitotic bookmarking by hematopoietic transcription factor GATA1. Kalmar, T. Regulated fluctuations in nanog expression mediate cell fate decisions in embryonic stem cells.

Kalverda, B. Characterization of genome-nucleoporin interactions in Drosophila links chromatin insulators to the nuclear pore complex. Cell Cycle 9, — Nucleoporins directly stimulate expression of developmental and cell-cycle genes inside the nucleoplasm.

Kaneko, S. Karachentsev, D. PR-Set7-dependent methylation of histone H4 Lys 20 functions in repression of gene expression and is essential for mitosis.

Karnani, N. Genomic study of replication initiation in human chromosomes reveals the influence of transcription regulation and chromatin structure on origin selection.

Cell 21, — Kehat, I. Modulation of chromatin position and gene expression by HDAC4 interaction with nucleoporins. Kemp, M. The histone deacetylase inhibitor trichostatin A alters the pattern of DNA replication origin activity in human cells. Kim, J. Integrative analysis of gene amplification in Drosophila follicle cells: parameters of origin activation and repression.

Kind, J. Single-cell dynamics of genome-nuclear lamina interactions. Kosak, S. Subnuclear compartmentalization of immunoglobulin loci during lymphocyte development. Koseoglu, M. Krajewski, W. A motif within SET-domain proteins binds single-stranded nucleic acids and transcribed and supercoiled DNAs and can interfere with assembly of nucleosomes.

Kueh, A. HBO1 is required for H3K14 acetylation and normal transcriptional activity during embryonic development. Kuilman, T. Identification of Cdk targets that control cytokinesis. Kundu, S. Kuzmichev, A. Histone methyltransferase activity associated with a human multiprotein complex containing the Enhancer of Zeste protein.

Lachner, M. Methylation of histone H3 lysine 9 creates a binding site for HP1 proteins. Laine, J. A physiological role for gene loops in yeast. LaJeunesse, D. E z : a polycomb group gene or a trithorax group gene? Development , — Lange, C. Cdks and cyclins link G1 length and differentiation of embryonic, neural and hematopoietic stem cells.

Lanzuolo, C. PcG complexes set the stage for epigenetic inheritance of gene silencing in early S phase before replication. Laurell, E. Phosphorylation of Nup98 by multiple kinases is crucial for NPC disassembly during mitotic entry.

Law, J. Establishing, maintaining and modifying DNA methylation patterns in plants and animals. Lee, S. Cell 43, — Lee, T. Control of developmental regulators by Polycomb in human embryonic stem cells. Lemaitre, J. Mitotic remodeling of the replicon and chromosome structure. Li, L. Coexistence of quiescent and active adult stem cells in mammals.

Li, V. Gap 1 phase length and mouse embryonic stem cell self-renewal. Li, Y. Liang, Y. Dynamic association of NUP98 with the human genome. Light, W. Z incorporation and INO1 transcriptional memory. Cell 40, — A conserved role for human Nup98 in altering chromatin structure and promoting epigenetic transcriptional memory.

Lipinszki, Z. Affinity purification of protein complexes from Drosophila embryos in cell cycle studies. Methods Mol. Liu, W. PHF8 mediates histone H4 lysine 20 demethylation events involved in cell cycle progression. Lo, S. A bridging model for persistence of a polycomb group protein complex through DNA replication in vitro. Cell 46, — Lubelsky, Y. DNA replication and transcription programs respond to the same chromatin cues. Luthra, R. Ma, T. MacAlpine, H. Drosophila ORC localizes to open chromatin and marks sites of cohesin complex loading.

Maeshima, K. Nuclear pore formation but not nuclear growth is governed by cyclin-dependent kinases Cdks during interphase. Cell-cycle-dependent dynamics of nuclear pores: pore-free islands and lamins.

Malhas, A. Defects in lamin B1 expression or processing affect interphase chromosome position and gene expression. Manansala, M. Margueron, R. Role of the polycomb protein EED in the propagation of repressive histone marks. Chromatin structure and the inheritance of epigenetic information. Martinez-Balbas, M. Displacement of sequence-specific transcription factors from mitotic chromatin.

Cell 83, 29— Marzluff, W. Metabolism and regulation of canonical histone mRNAs: life without a poly A tail. Mazouzi, A. DNA replication stress: causes, resolution and disease. McConnell, K.

Mechali, M. Eukaryotic DNA replication origins: many choices for appropriate answers. Mejlvang, J. New histone supply regulates replication fork speed and PCNA unloading.

Mesner, L. Bubble-chip analysis of human origin distributions demonstrates on a genomic scale significant clustering into zones and significant association with transcription. Miotto, B. HBO1 histone acetylase is a coactivator of the replication licensing factor Cdt1. Cell 37, 57— Moudry, P. Cell Death Differ. Muhlhausser, P. An in vitro nuclear disassembly system reveals a role for the RanGTPase system and microtubule-dependent steps in nuclear envelope breakdown.

Muller, J. Histone methyltransferase activity of a Drosophila Polycomb group repressor complex. Mummery, C. Cell cycle analysis during retinoic acid induced differentiation of a human embryonal carcinoma-derived cell line. Cell Differ. Nagai, S. Naumova, N. Organization of the mitotic chromosome. Nicolay, B. Combined inactivation of pRB and hippo pathways induces dedifferentiation in the Drosophila retina. Nishioka, K. PR-Set7 is a nucleosome-specific methyltransferase that modifies lysine 20 of histone H4 and is associated with silent chromatin.

Nizami, Z. The Cajal body and histone locus body. Cold Spring Harb. Nordman, J. Regulation of DNA replication during development. Oda, H. Monomethylation of histone H4-lysine 20 is involved in chromosome structure and stability and is essential for mouse development. Onischenko, E. Cdk1 and okadaic acid-sensitive phosphatases control assembly of nuclear pore complexes in Drosophila embryos. Cell 16, — Ono, T.

Differential contributions of condensin I and condensin II to mitotic chromosome architecture in vertebrate cells. Orkin, S. Chromatin connections to pluripotency and cellular reprogramming. Oza, P. Mechanisms that regulate localization of a DNA double-strand break to the nuclear periphery. Pajcini, K. Transient inactivation of Rb and ARF yields regenerative cells from postmitotic mammalian muscle.

Cell Stem Cell 7, — Pak, D. Association of the origin recognition complex with heterochromatin and HP1 in higher eukaryotes. Cell 91, — Pastor, W. Genome-wide mapping of 5-hydroxymethylcytosine in embryonic stem cells.

Pauklin, S. The cell-cycle state of stem cells determines cell fate propensity. Peric-Hupkes, D. Molecular maps of the reorganization of genome-nuclear lamina interactions during differentiation. Cell 38, — Perry, J. Petruk, S. Pope, B. Topologically associating domains are stable units of replication-timing regulation. Probst, A. Epigenetic inheritance during the cell cycle. Proffitt, J. Ramer, M. Dbf4 and Cdc7 proteins promote DNA replication through interactions with distinct Mcm protein subunits.

Reddy, K. Transcriptional repression mediated by repositioning of genes to the nuclear lamina. Rice, J. Mitotic-specific methylation of histone H4 Lys 20 follows increased PR-Set7 expression and its localization to mitotic chromosomes. Richmond, T. The cell grows and accumulates the building blocks of chromosomal DNA and the associated proteins as well as sufficient energy reserves to complete the task of replicating each chromosome in the nucleus.

The synthesis phase of interphase takes the longest because of the complexity of the genetic material being duplicated. Throughout interphase, nuclear DNA remains in a semi-condensed chromatin configuration.

In the S phase, DNA replication results in the formation of identical pairs of DNA molecules, sister chromatids, that are firmly attached to the centromeric region. The centrosome is duplicated during the S phase. The two centrosomes will give rise to the mitotic spindle, the apparatus that orchestrates the movement of chromosomes during mitosis. At the center of each animal cell, the centrosomes of animal cells are associated with a pair of rod-like objects, the centrioles, which are at right angles to each other.

Centrioles help organize cell division. Centrioles are not present in the centrosomes of other eukaryotic species, such as plants and most fungi. In the G 2 phase, the cell replenishes its energy stores and synthesizes proteins necessary for chromosome manipulation. Some cell organelles are duplicated, and the cytoskeleton is dismantled to provide resources for the mitotic phase. There may be additional cell growth during G 2. The final preparations for the mitotic phase must be completed before the cell is able to enter the first stage of mitosis.

During the multistep mitotic phase, the cell nucleus divides, and the cell components split into two identical daughter cells. The mitotic phase is a multistep process during which the duplicated chromosomes are aligned, separated, and move into two new, identical daughter cells. The first portion of the mitotic phase is called karyokinesis or nuclear division.

The second portion of the mitotic phase, called cytokinesis, is the physical separation of the cytoplasmic components into the two daughter cells. Karyokinesis, also known as mitosis, is divided into a series of phases prophase, prometaphase, metaphase, anaphase, and telophase that result in the division of the cell nucleus. Stages of the Cell Cycle : Karyokinesis or mitosis is divided into five stages: prophase, prometaphase, metaphase, anaphase, and telophase. The images at the bottom were taken by fluorescence microscopy hence, the black background of cells artificially stained by fluorescent dyes: blue fluorescence indicates DNA chromosomes and green fluorescence indicates microtubules spindle apparatus.

The membranous organelles such as the Golgi apparatus and endoplasmic reticulum fragment and disperse toward the periphery of the cell. The nucleolus disappears and the centrosomes begin to move to opposite poles of the cell. Microtubules that will eventually form the mitotic spindle extend between the centrosomes, pushing them farther apart as the microtubule fibers lengthen.

The sister chromatids begin to coil more tightly with the aid of condensin proteins and become visible under a light microscope. The remnants of the nuclear envelope fragment. The mitotic spindle continues to develop as more microtubules assemble and stretch across the length of the former nuclear area. Chromosomes become more condensed and discrete. Each sister chromatid develops a protein structure called a kinetochore in the centromeric region. The proteins of the kinetochore attract and bind mitotic spindle microtubules.

Kinetochore and Mitotic Spindle : During prometaphase, mitotic spindle microtubules from opposite poles attach to each sister chromatid at the kinetochore. In anaphase, the connection between the sister chromatids breaks down and the microtubules pull the chromosomes toward opposite poles. The exact length of the DNA segment associated with each histone core varies from species to species, but most such segments are approximately base pairs in length.

Furthermore, each histone molecule within the core particle has one end that sticks out from the particle. These ends are called N-terminal tails , and they play an important role in higher-order chromatin structure and gene expression. Figure 4: The nucleosome structure within chromatin.

Each nucleosome contains eight histone proteins blue , and DNA wraps around these histone structures to achieve a more condensed coiled form. Figure 5: To better fit within the cell, long pieces of double-stranded DNA are tightly packed into structures called chromosomes.

Although nucleosomes may look like extended "beads on a string" under an electron microscope, they appear differently in living cells. In such cells, nucleosomes stack up against one another in organized arrays with multiple levels of packing. The first level of packing is thought to produce a fiber about 30 nanometers nm wide. These 30 nm fibers then form a series of loops, which fold back on themselves for additional compacting Figure 5. The multiple levels of packing that exist within eukaryotic chromosomes not only permit a large amount of DNA to occupy a very small space, but they also serve several functional roles.

For example, the looping of nucleosome-containing fibers brings specific regions of chromatin together, thereby influencing gene expression. In fact, the organized packing of DNA is malleable and appears to be highly regulated in cells. Chromatin packing also offers an additional mechanism for controlling gene expression. Specifically, cells can control access to their DNA by modifying the structure of their chromatin. Highly compacted chromatin simply isn't accessible to the enzymes involved in DNA transcription , replication , or repair.

Thus, regions of chromatin where active transcription is taking place called euchromatin are less condensed than regions where transcription is inactive or is being actively inhibited or repressed called heterochromatin Figure 6. Figure 6: The structure of chromatin in interphase Heterochromatin is more condensed than euchromatin. Typically, the more condensed chromatin is, the less accessible it is by transcription factors and polymerases.

The dynamic nature of chromatin is regulated by enzymes. For example, chromatin can be loosened by changing the position of the DNA strands within a nucleosome. This loosening occurs because of chromatin remodeling enzymes, which function to slide nucleosomes along the DNA strand so that other enzymes can access the strand. This process is closely regulated and allows specific genes to be accessed in response to metabolic signals within the cell.

Another way cells control gene expression is by modifying their histones with small chemical groups, such as methyl and acetyl groups in the N-terminal tails that extend from the core particle. Different enzymes catalyze each kind of N-terminal modification. Scientists occasionally refer to the complex pattern of histone modification in cells as a "histone code.

In electron micrographs, eukaryotic interphase chromatin appears much like a plate of spaghetti — in other words, there is no obvious pattern of organization. In recent years, however, investigators have begun using fluorescent probes for each of the different interphase chromosomes.

In doing so, they have discovered that these chromosomes have functional and decidedly nonrandom arrangements. One of the first things these scientists noted was that uncondensed chromosomes occupy characteristic regions of the nucleus, which they termed chromosome territories. The spatial localization of these territories is thought to be important for gene expression.

In fact, with the advent of gene-specific probes, researchers are beginning to understand how the arrangement of chromosome territories can bring particular genes closer together. A second major observation related to chromosome territories is that the position of chromosomes relative to one another differs from cell to cell. Such differences reflect variation in gene expression patterns. This page appears in the following eBook.

Aa Aa Aa. What Are Chromosomes? How Are Eukaryotic Chromosomes Structured? Figure 3. Figure 6: The structure of chromatin in interphase.



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