Cell Nucleus

From CellBiology

Contents
1 Introduction
1.1 Lecture Audio
2 Objectives
3 Eukaryotes
4 Eukaryote Gene Expression
4.1 DNA -> mRNA -> Protein
4.2 Nucleus
4.3 Cytoplasm
5 Membrane Evolution
6 Nuclear Compartment
6.1 Nucleus Size
7 Nuclear Envelope
8 Nuclear Cytoskeleton
8.1 Nuclear Lamins Eukaryotic Cell Physical Compartments
9 Nuclear Transport History
10 Nuclear Pores
11 Nuclear Bodies
11.1 Cajal Bodies
11.2 PML Bodies
12 Chromosomes
12.1 Chromosome Territories
13 Nucleolus
13.1 Appearance
13.2 Function
14 Chromosome Features
14.1 Telomere
14.2 Centromere
14.3 Replication Origins
15 Chromosome DNA Packing
15.1 Nucleosomes
15.2 Histones
16 Abnormalities
16.1 Hutchinson-Gilford Progeria Syndrome
16.2 Emery-Dreifuss Muscular Dystrophy
17 Eukaryote Gene Expression
17.1 Protein Modification
17.2 Exocytosis
18 History
19 References
19.1 Textbooks
19.2 Books
19.3 Reviews
19.4 Articles
20 Acronyms
21 Movies
22 2016 Course Content

Introduction
This lecture introduces the nucleus and how information is transferred from stable stored information (DNA) converted to an intermediate
(mRNA, rRNA, tRNA) of variable stability, exported from the nucleus to the cytoplasm where mRNA is then translated into Protein. This is
gene expression, the products of this process are used either within the cell, exported (exocytosis) or used to replace worn out components.

We will study this topic looking at the key organelle in this process, the nucleus.

Nucleolar Assembly​ Movie | Movie - nuclear pore complexes (http://jcb.rupress.org/cgi/content/full/154/6/1147/FIG6/DC1)

Lecture Audio

Lecture Archive: 2015 (https://cellbiology.med.unsw.edu.au/cellbiology/index.php?title=Cell_Nucleus&oldid=51902) | 2014

(http://php.med.unsw.edu.au/cellbiology/index.php?title=Cell_Nucleus&oldid=48075) | 2013
(http://php.med.unsw.edu.au/cellbiology/index.php?title=Cell_Nucleus&oldid=42432) | 2010 | 2009 | 2008 Online content
(http://cellbiology.med.unsw.edu.au/units/science/lecture0804.htm)

MH - note that content listed below will not match exactly current lecture structure but has been selected as having similar content.

Objectives
Understand the concept of the cell nucleus
Understand the overall structure and components within the nucleus
Understand the functions of the nucleus
Brief understanding of chromosomal structure

Eukaryotes
Difference between Prokaryotes and Eukaryotes

Cytoskeleton
DNA structure
circular, linear
Packing (histones)
RNA processing (splicing)

Eukaryote Gene Expression

DNA -> mRNA -> Protein

DNA (transcription) -> mRNA (translation) -> Protein (function)

Nucleus
DNA (transcription) -> mRNA Nuclear processing (export)

DNA -> mRNA splicing (introns removed, exons joined) -> mRNA

Cytoplasm

mRNA (translation by ribosomes) -> Protein (processing)

rough endoplasmic reticulum (tem)
Protein Processing cytoplasm (free ribosomes), rough endoplasmic reticulum
(bound ribosomes)

Some proteins are returned to the nucleus by a Nuclear Localization Sequence (NLS)

SV40 Large T-antigen (http://www.plosone.org/article/fetchObject.action?

uri=info:doi/10.1371/journal.pone.0010475.g001&representation=PNG_M) - PKKKRKV (single part)
nucleoplasmin - KR[PAATKKAGQA]KKKK (two part) two clusters of basic amino acids separated by 10 amino acids.

Membrane Evolution
Postulated that an early "coating" structure lead to the infolding of the primitive plasma membrane to form the many membrane covered
organelles in the cytoplasm.

These modules may also be the evolutionary precursor to the nuclear pore structures and account for the double membrane that coats the
nucleus.

Nuclear Compartment
Nuclear envelope
Nuclear cytoskeleton
Nucleolus
Chromosome territories
 Euchromatin is a lightly packed form of chromatin (DNA, RNA and protein)
Heterochromatin is a tightly packed form of DNA
Interchromatin compartment
“speckles” interchromatin granule clusters
Splicing speckles or SC 35 domains
thought to be sites of storage of mRNA splicing factors
nuclear bodies - Cajal and PML

Links: MBOC - A cross-sectional view of a typical cell nucleus (http://www.ncbi.nlm.nih.gov/books/NBK26821/?

rendertype=figure&id=A606)

Nucleus Size
cell "karyoplasmic ratio" relatively constant (ratio of nuclear volume to cell volume)
most other cellular organelles (ER and mitochondria) can vary greatly in
amounts
multinucleated fission yeast cells
relative amount of cytoplasm surrounding each nucleus controls the size of
individual nuclei

Links: Nuclear size control in fission yeast. Neumann FR, Nurse P. J Cell Biol. 2007 Nov
19;179(4):593-600. PMID: 17998401 (http://www.ncbi.nlm.nih.gov/pubmed/17998401)

Nuclear Envelope
Forms structural compartment
Nuclear envelope two concentric membranes
Breaks down each mitosis (recycled)
Outer membrane continuous with Endoplasmic Reticulum (Endoplasmic Reticulum
is covered in Lecture 5)
Contains holes “nuclear pores” Nucleus Membrane Evolution

Nuclear envelope and endoplasmic reticulum system

Nuclear architecture
The Cell - Figure 8.1. The nuclear envelope (http://www.ncbi.nlm.nih.gov/books/bv.fcgi?
highlight=nuclear%20envelope&rid=cooper.figgrp.1325)

Nuclear Cytoskeleton

The nuclear cytoskeleton has 2 layers

outer - less organised surrounds nuclear envelope

inner - nuclear lamina - thin shell (20 nm) underlying the nuclear envelope
associates with both the inner nuclear membrane and underlying chromatin
can regulate gene expression
provides anchor sites for nuclear pore complexes
broken down each cell division

Cytoplasmic Nuclear pores (red) Nuclear Lamin A Lamin B1 in mitosis Nuclear lamins and
cytoskeleton migrating cytoskeleton (green) splicing factors
neuron chromatin (blue)
 Muscle satellite cell heterochromatin (green)
lamin A/C expression LaminA/C (red)[1]

Links: MBOC - A cross-sectional view of a typical cell nucleus (http://www.ncbi.nlm.nih.gov/books/NBK26821/?

rendertype=figure&id=A606)

Nuclear Lamins
Intermediate filaments covered in Cytoskeleton Lecture – Intermediate Filaments

Lamins - Class V intermediate filaments

Vertebrate lamins are classified into 2 types - A and B
10 nm in diameter, forms rope-like networks
polypeptide form dimers - central alpha-helical regions of two polypeptide chains
are wound around each other
assembly - head-to-tail association of dimers form linear polymers, side-by-side
association of polymers form filaments
B-type - B1 and B2 (586 aa protein, Mr 66,334 Da) lamins are ubiquitously
expressed throughout development
A-type - lamins A and C (Mr 74 kD and 65 kD) lamins in many organisms
expression does not appear until midway through embryogenesis (possible role in
differentiation)
lamins phosphorylation state affects nuclear envelope assembly state Nuclear Envelope showing underlying nuclear
(dephosphorylation nuclear envelope assembly, phosphorylation nuclear envelope matrix (green) (EM Image Martin Goldberg)
disassembly)
Lamins also link DNA to nuclear envelope (Lamin B1 interacts with chromatin)

Nuclear lamina and lamina-interacting Nucleoskeleton to the cytoskeleton

proteins complexes

Lamin Abnormalities - (laminopathies) mutations in lamins can lead to human disease (Hutchinson-Gilford Progeria Syndrome)

Links: MBOC - A cross-sectional view of a typical cell nucleus (http://www.ncbi.nlm.nih.gov/books/bv.fcgi?&rid=mboc4.figgrp.606) |

Nature Medicine - Image - The nuclear membranes include the interconnected but distinct inner and outer nuclear membranes and the nuclear
pore membrane (http://www.nature.com/nm/journal/v6/n2/fig_tab/nm0200_136_F1.html) MBOC - The breakdown and re-formation of the
nuclear envelope during mitosis (http://www.ncbi.nlm.nih.gov/books/bv.fcgi?&rid=mboc4.figgrp.2174) | The Cell - Intermediate Filament
Proteins (http://www.ncbi.nlm.nih.gov/books/bv.fcgi?&rid=cooper.table.1810) | The Cell - Model of lamin assembly
(http://www.ncbi.nlm.nih.gov/books/bv.fcgi?&rid=cooper.figgrp.1328) | Abcam - Lamin B1 antibody - Nuclear Envelope Marker
(http://www.abcam.com/index.html?datasheet=16048#ab16048-IF-Im1.jpg) | OMIM - Lamin A/C
(http://www.ncbi.nlm.nih.gov/entrez/dispomim.cgi?id=150330) | OMIM - Lamin B1 (http://www.ncbi.nlm.nih.gov/entrez/dispomim.cgi?
id=150340) |

Nuclear Transport History

1999 Nobel Prize Medicine - Günter Blobel for the discovery that "proteins have intrinsic signals that govern their transport and localization
in the cell"

Links: 1999 Nobel Prize Medicine - Günter Blobel (http://nobelprize.org/nobel_prizes/medicine/laureates/1999/press.html)

Nuclear Pores
 Protein complex
External diameter of about 120 nm (30 times the size of a ribosome)
Channel diameter 25 nm
channels between nucleus and cytoplasm (import/export)
passive passage of small polar molecules, ions, (<40-60 kDa)
active (selective/ regulated) passage of larger macromolecules, proteins and RNAs
importin - cytosolic protein

Nuclear pore complex

Nuclear pore EM structure timeline[2]

 Links: JCB Movie - Nuclear Pore Complex movement in interphase
(http://jcb.rupress.org/content/suppl/2001/07/11/200101089.F4.DC1)

Nuclear Bodies
Functional compartments

Nucleus Movie 1

Cajal Bodies

also called - nucleolar accessory bodies, coiled body, gems

0.1 - 2.0 microns, 1-10/ nucleus
Gems and Cajal bodies two forms of same structure
GEMS (Gemini of coiled bodies)

proposed sites where small nuclear ribonucleoproteins (snRNPs) and small nucleolar RNAs (snoRNPs) are modified.
snRNPs are particles that combine with pre-mRNA and various proteins to form spliceosomes
snoRNAs are a class of small RNA molecules that are involved with modifications of ribosomal RNAs (rRNAs) and other RNA
genes

Cajal bodies were first reported in 1903 by the Spanish cytologist/histologist Ramón y Cajal, who christened them "nucleolar accessory
bodies".

Cajal, S.R. 1903. Un sencillo método de coloración selectiva del retículo protoplásmico y sus efectos en los diversos órganos nerviosos de
vertebrados e invertebrados. Tra. Lab. Invest. Biol. 2:129–221.

See also Nature Reviews - The centennial of the Cajal body

PML Bodies

promyelocytic leukaemia nuclear bodies

also called PODs, ND10 or Kremer bodies
Function unknown
regulation of diverse cellular functions?
viral infection, cellular transformation, innate immunity, growth control, apoptosis
 dynamic hubs sensing stress and DNA damage

Chromosomes
not “visible” at interphase, condense for mitosis (1,000 fold)
condensation allows chromosomes to move along mitotic spindle without breaking or tangling
eukaryotes have separate chromosomes
Human 23 pairs, 22 autosome pairs, 2 sex chromosomes
diploid 2 copies of each chromosome (inherited one male/one female)
except male sex chromosomes X from mother Y from father
DNA and protein
packing of DNA
DNA structure
encodes genome (humans 30,000 genes, draft sequence published in 2001)
DNA genes encode RNA and proteins
DNA can also encodes nothing

Chromosome Territories
 Space within the nucleus occupied by individual chromosomes
Several different models as to how these territories interact
Intrachromosomal domains possibly RNA processing and transport

Links: MBOC - Selective painting of two interphase chromosomes in a human peripheral lymphocyte
(http://www.ncbi.nlm.nih.gov/books/bv.fcgi?&rid=mboc4.figgrp.686) | The Cell - Chromosome Territories
(http://www.ncbi.nlm.nih.gov/books/bv.fcgi?&rid=cooper.figgrp.1352)

Nucleolus
Appearance

Fibrillar center, dense fibrillar component, and granular component

Nucleolus changes during the cell cycle:
during mitosis - nucleolus breaks up as chromosomes condense
after mitosis - nucleolus reforms from coalesce of tips of 10 chromosomes

Function
Sites of ribosomal (rRNA) gene transcription, processing, and ribosome assembly
Nucleolus size depends on cell metabolic activity
Sites of ribosomal (rRNA) gene transcription, processing, and ribosome assembly
All cells contain multiple copies of rRNA genes

Links: Movie - Dynamics of nucleolus-derived foci throughout the cytoplasm and the disappearance of NDF during telophase
(http://jcb.rupress.org/content/suppl/2000/08/03/150.3.433.F2.DC1) | Movie - Dynamics of nucleolar reassembly in telophase were
analyzed by the visualization of fibrillarin-GFP (http://jcb.rupress.org/content/suppl/2000/08/03/150.3.433.F3.DC1)

Chromosome Features

2 telomeres, centromere, replication origins

Telomere- at ends of chromosome (bacterial DNA circular)
Centromere- holds duplicated DNA together
Kinetochore - protein complex forms around the centromere forms during mitosis
Chromatin - DNA packed by DNA binding proteins (histones and non-histones) form 30nm DNA fibre
2 types of chromatin in interphase nuclei (based on cytology)
heterochromatin - highly condensed (restricted gene transcription)
euchromatin - less condensed (gene transcription)

Telomere
 at ends of all chromosomes (not bacterial DNA circular)
roles in chromosome replication and maintenance
replication
for replicating the ends of linear chromosomes
maintenance
proposed to provide each cell with a replication counting mechanism that helps prevent unlimited proliferation
each cell division shortens telomere 50–100 nucleotides
DNA 100s to 1,000s repeats of a simple-sequence containing clusters of G residues (humans AGGGTT)
Telomerase enzyme maintains length

Centromere

directs movement of each chromosome into daughter cells every time a cell divides
centromere embedded in heterochromatin
satellite DNA sequences (AT-rich) repeated many thousands of times
proteins assemble on this to form Kinetochore
attachment site for spindle microtubules

Links: MBOC - Centromere (http://www.ncbi.nlm.nih.gov/books/bv.fcgi?&rid=mboc4.figgrp.672)

Replication Origins
 DNA replication initiates at multiple origins (ori)
in both prokaryotic and eukaryotic DNA
multiple origins in eukaryotes (human genome about 30,000 origins)
each origin produces two replication forks (moving in opposite directions)

Chromosome DNA Packing

Euchromatin ("good chromatin") - light, transcriptionally active, about 10% of all

chromatin.

Heterochromatin - condensed, transcriptionally inactive, about 90% of all chromatin.

Nucleosomes
formed by DNA wrapped around histones
unit particle of chromatin (nucleosomal histones) (discovered 1974)
EM unfolded DNA has "beads on a string" appearance
second order folding forms 300 nm fibre
Chromosome-condensation
condensed DNA for mitosis 700 nm fibre

Histones

only in eukaryotes
small proteins positively charged (binds negatively charged DNA)
not sequence specific binding (as in transcription factors)
4 core histones (H2A, H2B, H3, and H4)
2 linker histones (H1/H5)

Abnormalities
Hutchinson-Gilford Progeria Syndrome
In more than 80% of cases the gene defect responsible for HGPS is a single spontaneous mutation in codon 608 of the LMNA gene,
which encodes both lamin A and lamin C
 single-base substitution in exon 11 that reveals a cryptic splice site in the LMNA
gene thus producing a truncated protein
progerin is a mutant form of the nuclear architectural protein lamin A

Emery-Dreifuss Muscular Dystrophy

OMIM - LAMIN A/C; LMNA (http://www.ncbi.nlm.nih.gov/entrez/dispomim.cgi?
id=150330)

Loss of a-Type Lamin Expression Compromises Nuclear Envelope Integrity Leading to

Muscular Dystrophy (http://jcb.rupress.org/cgi/content/abstract/147/5/913)

Eukaryote Gene Expression

Nucleosome
DNA -> mRNA -> Protein

DNA (transcription) -> mRNA (translation) -> Protein (function)

DNA -> mRNA splicing (introns removed, exons joined) -> mRNA

DNA -> rRNA, tRNA, siRNA (RNA interference (RNAi) pathway

Nucleus DNA (transcription) -> mRNA Nuclear processing (export) Cytoplasm mRNA
(translation) -> Protein (cytoplasm, rough endoplasmic reticulum)

Protein Modification

Protein - cytoplasmic (free ribosomes), rough endoplasmic reticulum (bound ribosomes)

Hutchinson-Gilford_Progeria_Syndrome
Exocytosis

Rough Endoplasmic Reticulum -> transport vesicle -> Golgi apparatus -> secretory
vesicle

History
Below are some example historical research finding related to cell junctions from the JCB
Archive (http://jcb.rupress.org/misc/fromthearchive.shtml) and other sources.

1961 The nucleolar origin of rRNA (http://jcb.rupress.org/cgi/content/full/168/4/524-a)

Base compositions and half-lives suggest to Jan-Erik Edström that the nucleolus is the
source of rRNA.

References
1. Neveen A Hosny, Mingying Song, John T Connelly, Simon Ameer-Beg, Martin M
Knight, Ann P Wheeler Super-resolution imaging strategies for cell biologists
using a spinning disk microscope. PLoS ONE: 2013, 8(10);e74604 PubMed
24130668 | PLoS One.
(http://www.plosone.org/article/info:doi/10.1371/journal.pone.0074604)
2. Alexander von Appen, Martin Beck Structure Determination of the Nuclear Pore
Complex with Three-Dimensional Cryo electron Microscopy. J. Mol. Biol.:
2016; PubMed 26791760

Textbooks

Molecular Biology of the Cell (4th ed.) - Part 1 Chapter 8 - The Cell Nucleus -
Three views of a cell membrane (http://www.ncbi.nlm.nih.gov:80/books/bv.fcgi?
db=Books&rid=mboc4.figgrp.1862) Lamin A EM - normal and KO
Molecular Cell Biology - The Dynamic Cell
(http://www.ncbi.nlm.nih.gov:80/books/bv.fcgi?db=Books&rid=mcb.chapter.145)
The Cell- A Molecular Approach - An Overview of Cells and Cell Research (http://www.ncbi.nlm.nih.gov:80/books/bv.fcgi?
db=Books&rid=cooper.chapter.89)
Search Online Textbooks

"cell nucleus" Molecular Biology of the Cell (http://www.ncbi.nlm.nih.gov:80/entrez/query.fcgi?

db=Books&cmd=search&doptcmdl=DocSum&term=cell+nucleus+AND+mboc4%5Bbook%5D) | Molecular Cell Biology
(http://www.ncbi.nlm.nih.gov:80/entrez/query.fcgi?
db=Books&cmd=search&doptcmdl=DocSum&term=cell+nucleus+AND+mcb%5Bbook%5D) | The Cell- A molecular Approach
(http://www.ncbi.nlm.nih.gov:80/entrez/query.fcgi?
db=Books&cmd=search&doptcmdl=DocSum&term=cell+nucleus+AND+cooper%5Bbook%5D)
"nuclear envelope" Molecular Biology of the Cell (http://www.ncbi.nlm.nih.gov:80/entrez/query.fcgi?
db=Books&cmd=search&doptcmdl=DocSum&term=nuclear+envelope+AND+mboc4%5Bbook%5D) | Molecular Cell Biology
(http://www.ncbi.nlm.nih.gov:80/entrez/query.fcgi?
db=Books&cmd=search&doptcmdl=DocSum&term=nuclear+envelope+AND+mcb%5Bbook%5D) | The Cell- A molecular Approach
(http://www.ncbi.nlm.nih.gov:80/entrez/query.fcgi?
db=Books&cmd=search&doptcmdl=DocSum&term=nuclear+envelope+AND+cooper%5Bbook%5D)
"nuclear pore" Molecular Biology of the Cell (http://www.ncbi.nlm.nih.gov:80/entrez/query.fcgi?
db=Books&cmd=search&doptcmdl=DocSum&term=nuclear+pore+AND+mboc4%5Bbook%5D) | Molecular Cell Biology
(http://www.ncbi.nlm.nih.gov:80/entrez/query.fcgi?
db=Books&cmd=search&doptcmdl=DocSum&term=nuclear+pore+AND+mcb%5Bbook%5D) | The Cell- A molecular Approach
(http://www.ncbi.nlm.nih.gov:80/entrez/query.fcgi?
db=Books&cmd=search&doptcmdl=DocSum&term=nuclear+pore+AND+cooper%5Bbook%5D)

Books
The Cytoskeleton - cellular architecture and choreography (http://books.google.com/books?id=JNHuxHzTm7IC)

Reviews

Selma Osmanagic-Myers, Thomas Dechat, Roland Foisner Lamins at the crossroads of mechanosignaling. Genes Dev.: 2015, 29(3);225-37
PubMed 25644599

Katherine L Wilson, Scott C Dawson Evolution: functional evolution of nuclear structure. J. Cell Biol.: 2011, 195(2);171-81 PubMed
22006947

Michael D Huber, Larry Gerace The size-wise nucleus: nuclear volume control in eukaryotes. J. Cell Biol.: 2007, 179(4);583-4 PubMed
17998404

Roderick Y H Lim, Ueli Aebi, Birthe Fahrenkrog Towards reconciling structure and function in the nuclear pore complex. Histochem.
Cell Biol.: 2008, 129(2);105-16 PubMed 18228033

Articles
Wenjing Li, Brian J Ferguson, Walid T Khaled, Maxine Tevendale, John Stingl, Valeria Poli, Tina Rich, Paolo Salomoni, Christine J Watson
PML depletion disrupts normal mammary gland development and skews the composition of the mammary luminal cell progenitor
pool. Proc. Natl. Acad. Sci. U.S.A.: 2009, 106(12);4725-30 PubMed 19261859

J Ellenberg, E D Siggia, J E Moreira, C L Smith, J F Presley, H J Worman, J Lippincott-Schwartz Nuclear membrane dynamics and
reassembly in living cells: targeting of an inner nuclear membrane protein in interphase and mitosis. J. Cell Biol.: 1997, 138(6);1193-
206 PubMed 9298976

L Yang, T Guan, L Gerace Integral membrane proteins of the nuclear envelope are dispersed throughout the endoplasmic reticulum
during mitosis. J. Cell Biol.: 1997, 137(6);1199-210 PubMed 9182656

Acronyms
AA - amino acid
DNA - deoxyribonucleic acid
EM - electron microscopy
FL - flourescent
GEMS - Gemini of coiled bodies
INM - inner nuclear membrane
KASH - Klarsicht/ANC-1/Syne-1 homology domain–containing protein
LEM domain - fold identified in LAP2, emerin, and MAN1 confers direct binding to dsDNA
LINC - nucleoskeleton to the cytoskeleton complexes
mRNA - messenger RNA
NE - nuclear envelope
 NES - nuclear export signal
NLS - nuclear localization signal
NPC - nuclear pore complex
NUP - nucleoporin
ONM - outer nuclear membrane
OMIM - Online Mendelian Inheritance in Man Database
PML Bodies - promyelocytic leukaemia nuclear bodies
RER - rough endoplasmic reticulum
RNA - ribonucleic acid
rRNA - ribosomal RNA
SER - smooth endoplasmic reticulum
SUMO - small ubiquitin-related modifier
SUN - Sad1/UNC-84
tRNA - transfer RNA

Movies
Nuclear pore complexes are fixed in place Daigle et al. (http://jcb.rupress.org/cgi/content/abstract/154/1/71) report that nuclear pore
complexes (NPCs) undergo limited movements (http://jcb.rupress.org/cgi/content/full/200101089/F4/DC1) that match the deformations
of the nuclear envelope as tracked using a grid (http://jcb.rupress.org/cgi/content/full/200101089/F4/DC2) of bleached nuclear lamins.
NPCs are therefore remarkably stable complexes, and are probably anchored to a protein network in the nuclear envelope.

Nucleoporins reassemble around post-mitotic chromatin A conserved nuclear pore subcomplex was characterized and tracked by
Belgareh et al. (http://jcb.rupress.org/cgi/content/abstract/154/6/1147), who found that the proteins were recruited
(http://jcb.rupress.org/cgi/content/full/154/6/1147/FIG6/DC1) during telophase in a rim pattern surrounding the chromosomes. A low
level of staining was also apparent on the kinetochores throughout mitosis.

Nucleolar re-formation after mitosis Savino et al. (http://jcb.rupress.org/cgi/content/abstract/153/5/1097) follow the re-formation
(http://jcb.rupress.org/cgi/content/full/153/5/1097/F3/DC1) of nucleoli after mitosis. Prenucleolar bodies (PNB) form on the
chromosome surface and nucleolar material flows along links between PNBs
(http://jcb.rupress.org/cgi/content/full/153/5/1097/F4/DC2) and towards (http://jcb.rupress.org/cgi/content/full/153/5/1097/F6/DC1) a
developing nucleolar organizer region (http://jcb.rupress.org/cgi/content/full/153/5/1097/F4/DC1) (NOR). Eventually this leads to the
fusion (http://jcb.rupress.org/cgi/content/full/153/5/1097/F9/DC1) of nucleoli to form a single entity.

Processing complexes may help reassemble nucleoli Nucleolar reassembly (http://jcb.rupress.org/cgi/content/full/150/3/433/F3/DC2)

during telophase is shown by Dundr et al. (http://jcb.rupress.org/cgi/content/abstract/150/3/433) to require mitotically preserved
processing complexes.

Speckles - A splicing factor has limited mobility Based on the limited mobility
(http://jcb.rupress.org/cgi/content/full/150/1/41/F1/DC1) of a splicing factor, Kruhlak et al.
(http://jcb.rupress.org/cgi/content/abstract/150/1/41) determine that the factor undergoes frequent but transient interactions with
relatively immobile nuclear binding sites, both when associated with speckles and when dispersed in the nucleoplasm. This a 3-D video
that should be viewed using red/green 3-D glasses.

2016 Course Content

Lectures: Cell Biology Introduction | Cells Eukaryotes and Prokaryotes | Cell Membranes and Compartments | Cell
Nucleus | Cell Export - Exocytosis | Cell Import - Endocytosis | Cytoskeleton Introduction | Cytoskeleton -
Microfilaments | Cytoskeleton - Microtubules | Cytoskeleton - Intermediate Filaments | Cell Mitochondria | Cell Junctions
| Extracellular Matrix 1 | Extracellular Matrix 2 | Cell Cycle | Cell Division | Cell Death 1 | Cell Death 2 | Signal 1 | Signal
2 | Stem Cells 1 | Stem Cells 2 | Development | 2016 Revision Moodle
(http://moodle.telt.
id=18930)
Laboratories: Introduction to Lab | Microscopy Methods | Preparation/Fixation | Cell Knockout Methods | Cytoskeleton
Exercise | Immunochemistry | Project Work | Confocal Microscopy | Tissue Culture | Stem Cells Lab | Microarray Visit

2016 Projects:

Dr Mark Hill 2015, UNSW Cell Biology - UNSW CRICOS Provider Code No. 00098G

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