Cellular Biology

Part 5
The nucleus
5.1
The nucleus - Structure
Figure: A cell from the human cervical carcinoma cell line HeLa has a nucleus
that is easily seen using light microscopy.
Major compartments
1. Nuclear envelope
2. Nucleoplasm
3. Nucleolus
4. Nuclear Lamina
5. Chromatin
Fig. Nucleus, image copyright Sagar Aryal

1. The nuclear envelope
-Double membranes
- The outer nuclear membrane is continuous with

the membranes of the ER, and the lumen of the
nuclear envelope is continuous with the lumen of
the ER.
-The nuclear envelope contains numerous NPCs
Figure 09.F12: The nuclear

envelope is continuous with the
endoplasmic reticulum.
1. The nuclear envelope - NPCs
Large molecules are actively transported between the nucleus and cytoplasm
Uncharged molecules < 100 Da: pass through the nuclear envelope.
> 100 Da: NPCs
Particles up to 9 nm in diameter
(corresponding to globular proteins up to
40 kDa) can pass through NPCs by passive
diffusion, as can metabolites, nucleotides,
and other small molecules
The nuclear pore complex.

(Modified with permission from Alberts B,
Bray D, Lewis J, et al.: Molecular Biology
of the Cell, 3rd ed. New York, Garland Publishing, 1994.
The nuclear pore is made of about 30-100
nucleoporin proteins.
Many different classes of molecules and macromolecules are

transported through NPCs. Not shown are small, uncharged
molecules (<100 Da) that can diffuse through the membranes of
the nuclear envelope
Proteins are selectively transported into the nucleus through NPCs
Cores do not contain information for

nuclear entry
nuclear pore is a selective channel that

allows only proteins with the proper
information in their amino acid sequence
to enter
Figure: Injection of nucleoplasmin into the cytosol or

the nucleus of Xenopus oocytes shows the importance
of the C-terminal tail fragment.
Mature nuclear proteins contain sequence information required for their nuclear localization.
The signal within a protein that targets it to the nucleus is a stretch of amino acids termed
the NLS (nuclear localization sequence )
Export of proteins from the nucleus is also receptor-mediated
Short stretches of amino acids rich in leucine act as the most common nuclear
export sequences (NESs)
A nuclear export receptor binds

proteins that contain NESs in the
nucleus and transports them to the
cytoplasm
The NESs were discovered in the

course of studies of the growth of
human HIV in infected cells
Figure: Some proteins, such as HIV Rev, shuttle

between the nucleus and the cytoplasm.
Transport of most RNAs is unidirectional from the nucleus to the cytoplasm
2. The Nucleoplasm
- gelatinous substance within the nuclear envelope
- composed mainly of water with dissolved salts, enzymes, and organic molecules
- supports the nucleus by helping to maintain its shape

3. The Nucleolus
 a dense, membrane-less structure within the nucleus
 Genes that encode rRNAs are present on multiple chromosomes that cluster
together to form nucleolar subcompartments.
 rRNA is synthesized and ribosomal subunits are assembled in the nucleolus.
 disappears when a cell undergoes

division and is reformed after the
completion of cell division

4. The Nuclear Lamina
 Located beneath the inner nuclear membrane and is physically connected
to it by lamina-associated integral membrane proteins.
 Plays a role in nuclear envelope assembly and may provide physical support
for the nuclear envelope
 Protein complexes that interact with the

nuclear lamina cross the nuclear envelope
and link the cytoskeleton to the nuclear
interior
Yeast and some other unicellular

eukaryotes lack a nuclear lamina
Figure “ The nuclear lamina is anchored to

the inner nuclear membrane
5. The Chromatin
 consist of DNA, which contains heredity information and instructions for cell
growth, development, and reproduction
- Heterochromatin: highly condensed, transcriptionally inactive form

- Euchromatin: less densely compacted
Figure: Individual chromosomes

occupy distinct areas of the nucleus
called chromosome territories.
What is the advantage to a eukaryotic cell of having a nucleus?
Having a nucleus allows a cell to have

much more sophisticated regulation of
gene expression than is possible in
prokaryotic cells
The nucleus protects the DNA of the

cell
Figure: In prokaryotes, transcription and translation

are coupled (left). In eukaryotes, transcription and
translation occur in separate compartments (right)
Nuclei vary in appearance according to cell type and organism
 Nuclei range in size from about one micron (1 μm) to more than 10 μm in diameter
Most cells have a single nucleus, but some cells contain multiple nuclei, and a few cell
types lack nuclei
Figure: A Drosophila embryo at the

multinucleate stage
Nucleus origin
The nucleus may have arisen by endosymbiosis, a process in which one prokaryotic cell
engulfs another cell, which then becomes a primitive nucleus
5.2
Chromatin – chromosomes
Bacteria- genetic material: in the form of a nucleoid
Eukaryotic- genetic material: the mass of chromatin within the nucleus
The centromere is the specialized DNA sequence of a

chromosome that links a pair of sister chromatids
The nucleosome is the subunit of all chromatin
• Micrococcal nuclease releases individual nucleosomes from chromatin as ~10-

nm particles.
• A nucleosome contains ~200 bp of DNA, two copies of each core histone (H2A,
H2B, H3, and H4), and one copy of H1.
• DNA is wrapped around the outside surface of the protein octamer.
The nucleosome consists of approximately Sequences on the DNA that tie on different
equal masses of DNA and histones.. turns around the nucleosome may close
together.
A linker histone helps to pull
nucleosomes together and
pack them into a more
compact chromatin fiber.
Different levels of DNA packing
The 30 nm solenoid structure fiber

forms loops whose base is attached
to scaffold proteins.
Decondensation of these loops
allows the transcription of genes
they encode.
Chromosomes have banding patterns
• Certain staining techniques cause the

chromosomes to have the appearance of a
series of striations called G-bands.
• The bands are lower in G  C content than
the interbands.
• Genes are concentrated in the G  C-rich
interbands.
Figure: Every chromosome has a

distinct G-banding pattern.
Telomeres are the ends of chromosomes
 Protect ends
 Maintain length
Telomeres are made of repeating sequences of TTAGGG on one strand paired

with AATCCC on the other strand
Telomerase
It is required for cells that must divide many times
Cancer
Two closely related species can have similar genome sizes
but very different chromosome numbers
The 23 chromosome pairs of the human genome
FISH: fluorescent in situ hybridization Karyotype: chromosomes articially lined

up in order (e.g. for cytogenetic analysis)
X chromosomes undergo global changes
One of the two X chromosomes is inactivated at random in each cell during

embryogenesis of eutherian mammals
Dosage compensation: equalizes the level of expression of X-linked genes in

the two sexes
Different means of dosage compensation are used to equalize X chromosome expression

in male and female.
One of the two X chromosomes is
inactivated in the cells of mammalian
females by heterochromatin formation.
Each female cell contains two X
chromosomes, one from the mother (Xm)
and the other from the father (Xp).

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Part 5

Fig. Nucleus, image copyright Sagar Aryal

- The outer nuclear membrane is continuous with

-The nuclear envelope contains numerous NPCs

Figure 09.F12: The nuclear

The nuclear pore complex.

Many different classes of molecules and macromolecules are

Cores do not contain information for

nuclear pore is a selective channel that

Figure: Injection of nucleoplasmin into the cytosol or

A nuclear export receptor binds

The NESs were discovered in the

Figure: Some proteins, such as HIV Rev, shuttle

- gelatinous substance within the nuclear envelope

- supports the nucleus by helping to maintain its shape

Fig. Nucleus, image copyright Sagar Aryal

 a dense, membrane-less structure within the nucleus

 rRNA is synthesized and ribosomal subunits are assembled in the nucleolus.

 disappears when a cell undergoes

Fig. Nucleus, image copyright Sagar Aryal

 Protein complexes that interact with the

Yeast and some other unicellular

Figure “ The nuclear lamina is anchored to

- Heterochromatin: highly condensed, transcriptionally inactive form

Figure: Individual chromosomes

Having a nucleus allows a cell to have

The nucleus protects the DNA of the

Figure: In prokaryotes, transcription and translation

Figure: A Drosophila embryo at the

Eukaryotic- genetic material: the mass of chromatin within the nucleus

The centromere is the specialized DNA sequence of a

• Micrococcal nuclease releases individual nucleosomes from chromatin as ~10-

The 30 nm solenoid structure fiber

• Certain staining techniques cause the

Figure: Every chromosome has a

Telomeres are made of repeating sequences of TTAGGG on one strand paired

It is required for cells that must divide many times

FISH: fluorescent in situ hybridization Karyotype: chromosomes articially lined

One of the two X chromosomes is inactivated at random in each cell during

Dosage compensation: equalizes the level of expression of X-linked genes in

Different means of dosage compensation are used to equalize X chromosome expression

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