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Nucleolus

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The nucleolus is the largest structure found inside the nucleus of eukaryotic cells.

Diagram human cell nucleus multilang
Diagram human cell nucleus multilang

It serves as the primary site for ribosome biogenesis, which is the process of building ribosomes. Ribosomes are essential because they allow cells to perform vital work. The nucleolus is composed of DNA, RNA, and proteins. It forms around specific chromosomal areas known as nucleolar organizing regions, or NORs. Because it manages such critical tasks, the nucleolus is essential for a cell to function and survive.

Building ribosomes is a complex, multi-step biological process. First, the cell must perform transcription to create ribosomal RNA, or rRNA. This begins in the fibrillar center (FC), where rDNA transcription occurs. Next, the process moves to the dense fibrillar component (DFC). This area contains a protein called fibrillarin, which is necessary for rRNA processing. Finally, the granular component (GC) uses the protein nucleophosmin to assist in ribosome biogenesis.

NucleolusNCc
NucleolusNCc

This organized structure allows the cell to manufacture its tools efficiently.

In humans, the assembly of these ribosomes requires several different types of RNA polymerase. RNA polymerase I transcribes most rRNA transcripts, including the 28S, 18S, and 5.8S molecules. These are initially transcribed as a single, long precursor molecule called 45S pre-rRNA. To finish the job, the cell must remove internal and external spacers from this molecule. Meanwhile, RNA polymerase III transcribes the 5S rRNA subunit. In higher eukaryotes, this 5S rRNA is made in the nucleoplasm before traveling into the nucleolus to join the assembly.

NucleolusNCc
NucleolusNCc

To complete the ribosome, the rRNA must join with ribosomal proteins. These proteins are created through a different pathway in the nucleoplasm. The genes for these proteins are transcribed by RNA polymerase II. Once the proteins are fully made, they are imported into the nucleus and then into the nucleolus. Inside the nucleolus, the rRNA and the proteins associate to form two distinct parts. These are the 40S small subunit and the 60S large subunit. These subunits are then exported through nuclear pore complexes into the cytoplasm.

NucleolusNCc
NucleolusNCc

Scientists have been observing the nucleolus for nearly two centuries. It was first identified using bright-field microscopy during the 1830s. In 1839, Theodor Schwann wrote about small corpuscles in nuclei, calling them "Kernkörperchen." The English term "nucleolus" appeared in a 1947 translation of his work. A major breakthrough occurred in 1964 through the work of John Gurdon and Donald Brown. They studied the African clawed frog, *Xenopus laevis*, and discovered that eggs without a nucleolus could not survive. This proved that the nucleolus performs a function necessary for life.

There are many unique variations in how nucleoli appear and function. While most cells only show one or two, a single diploid human cell has ten nucleolus organizer regions. This means a cell has the potential to form more nucleoli. In many plant species, the nucleolus contains a clear central area called a nucleolar vacuole. Plant nucleoli also contain very high concentrations of iron compared to human or animal nucleoli.

NucleolusNCc
NucleolusNCc

To study these details, scientists use electron microscopes to see the ultrastructure or fluorescent protein tagging to watch how the parts move.

Beyond building ribosomes, the nucleolus also acts as a regulatory center. It can capture and immobilize certain proteins in a process called nucleolar detention. When proteins are detained, they cannot move or interact with their usual binding partners. This detention is managed by long noncoding RNAs that come from the intergenic regions of the nucleolus. This mechanism helps the cell respond to various types of stress. Because of this, malfunctions in the nucleolus can lead to human conditions called nucleolopathies. Scientists are currently investigating the nucleolus as a target for cancer chemotherapy.

NucleolusNCc
NucleolusNCc
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File:Diagram human cell nucleus multilang.svg
Diagram human cell nucleus multilang.svg
Widespread-Expression-of-BORISCTCFL-in-Nor...
File:NucleolusNCc.jpg
NucleolusNCc.jpg
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