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The Golgi apparatus transports and modifies proteins in eukaryotic cells. How have scientists studied dynamic protein
movements through the Golgi?
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The Golgi apparatus is the central organelle mediating protein and lipid transport within the eukaryotic cell. Typically textbooks illustrate the Golgi as something resembling a stack of pita bread.
However, this depiction does not adequately illustrate the dynamic nature of the Golgi compartments (called cisternae) or the variety of morphologies the Golgi manifests in different cell types. We
can learn a lot by simply asking why these diverse structures even exist. Researchers do not yet fully understand how various Golgi morphologies affect its function. However, scientists are
currently using the subtle variations in Golgi morphology among different cell types to ask how proteins move through the Golgi apparatus.
Each cisterna or region of the Golgi contains different protein modification enzymes. What do these enzymes
do? The Golgi enzymes catalyze the addition or removal of sugars from cargo proteins (glycosylation), the
addition of sulfate groups (sulfation), and the addition of phosphate groups (phosphorylation). Cargo proteins
are modified by enzymes (called resident enzymes) located within each cisterna. The enzymes sequentially
add the appropriate modifications to the cargo proteins. Some Golgi-mediated modifications act as signals to
direct the proteins to their final destinations within cells, including the lysosome and the plasma membrane.
What happens when there are defects in Golgi function? Defects in various aspects of Golgi function can
result in congenital glycosylation disorders, some forms of muscular dystrophy, and may contribute to
diabetes, cancer, and cystic fibrosis (Ungar 2009).
How do cargo proteins move between the Golgi cisternae? Scientists have proposed two possible
explanations: the vesicular transport model and cisternal maturation model. Interestingly, both models
account for the Golgi's steady state conditions and processes, yet they do so quite differently (Figure 2). In
2002 James Rothman and Randy Schekman won the Lasker Prize for their groundbreaking work detailing
Figure 1: The Golgi apparatus modifies and sorts proteins for transport
the membrane and vesicle systems that make secretion possible in eukaryotic cells. These two scientists
throughout the cell.
worked independently using different model organisms and different biological approaches (Strauss 2009).
The Golgi apparatus is often found in close proximity to the ER in cells.
Together they delivered strong evidence that there are common molecules and processes involved in
Protein cargo moves from the ER to the Golgi, is modified within the Golgi,
membrane fusion and fission in eukaryotes. Rothman and his colleagues biochemically reconstituted
and is then sent to various destinations in the cell, including the lysosomes
mammalian Golgi membranes, isolating vesicles capable of moving from one cisterna to another. As a
and the cell surface.
different approach, Schekman and his colleagues used yeast genetics to identify and characterize many of
© 2009 Nature Publishing Group Xu, D. & Esko, J. D. A Golgi-on-a-chip for
the important proteins involved in secretion in this single-celled eukaryote. Over time Rothman and
glycan synthesis. Nature Chemical Biology 5, 612–613 (2009). All rights
Schekman's work converged on several important molecules that were involved in vesicle formation and
reserved.
fusion, thus leading to what came to be called the vesicular transport model.
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Figure Detail
What were all the vesicles Rothman discovered doing in the Golgi? The current cisternal maturation model proposes that these vesicles are transport vehicles for Golgi enzymes rather than for
protein cargo. Retrograde vesicles that travel backward through the Golgi bud off of a cisterna to transfer enzymes to younger cisternae. Meanwhile other vesicles, arriving from older cisternae,
carry the enzymes necessary for the next steps in protein modification (Glick & Malhotra 1998; Pellham 1998).
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cisternae over time. The vesicular transport model would predict that an individual cis cisterna would remain cis, with Video courtesy of Dr. Benjamin S. Glick, University of
characteristic cis enzymes, over its entire lifespan. However, the cisternal maturation model would predict that a newly Chicago. All rights reserved.
formed cis cisterna would eventually mature into a medial, then a trans cisterna, before breaking apart when its contents
were packaged for their final destinations in the cell. In their experiments the two research groups linked fluorescent
proteins (glowing green or red) to the proteins present in different, individual cisternae of S. cerevisiae, and they followed these colored molecules over time. The researchers designed their
experiments to test the predictions of the vesicular transport and cisternal maturation models. If the vesicular transport model were correct, then the cisternae would be stable and maintain the
same fluorescently labeled Golgi resident proteins over time. In contrast, if the cisternal maturation model was not correct, then each cisterna would contain a changing set of Golgi proteins over
time. In their experiments, the researchers created beautiful movies of the yeast and observed that the individual cisternae changed color over time. After analyzing a variety of Golgi proteins, the
researchers consistently observed changes in the protein composition of individual cisternae over time. Their results provided strong evidence for the cisternal maturation model.
While some aspects of protein transport through the Golgi are better understood than they used to be, there are still many unresolved issues surrounding the specifics within different organisms.
Moreover, questions remain about the unifying characteristics that are shared between all Golgi. A recent gathering of prominent Golgi researchers identified several important questions to be
addressed in the future, including:
Do different types of secretory cargo follow distinct routes through the Golgi?
What molecular mechanisms drive and regulate cisternal maturation?
Are there specialized domains in the Golgi cisternae? How are they created, and what roles do they play in cargo sorting and export?
How are the Golgi compartments constructed and remodeled?
Is Golgi stacking fundamentally important for membrane traffic? If so, how do organisms such as S. cerevisiae bypass this requirement?" (Emr et al. 2009)
Summary
The structure of the Golgi apparatus varies in different cell types. The dispersed nature of Golgi cisternae in the yeast Saccharomyces cerevisiae allowed researchers to resolve individual
cisternae. By observing fluorescently labeled proteins that normal reside within different cisternae, researchers found convincing evidence that the Golgi cisternae change over time, supporting the
cisternal maturation model of protein movement through the Golgi apparatus. However, there is clearly much left to discover about the Golgi.
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