@article {5149,
	title = {The structure of purified kinetochores reveals multiple microtubule-attachment sites.},
	journal = {Nature Structural \& Molecular Biology},
	volume = {19},
	year = {2012},
	month = {2012 Aug 12},
	pages = {925-9},
	abstract = {<p>Chromosomes must be accurately partitioned to daughter cells to prevent aneuploidy, a hallmark of many tumors and birth defects. Kinetochores are the macromolecular machines that segregate chromosomes by maintaining load-bearing attachments to the dynamic tips of microtubules. Here, we present the structure of isolated budding-yeast kinetochore particles, as visualized by EM and electron tomography of negatively stained preparations. The kinetochore appears as an \&nbsp;126-nm particle containing a large central hub surrounded by multiple outer globular domains. In the presence of microtubules, some particles also have a ring that encircles the microtubule. Our data, showing that kinetochores bind to microtubules via multivalent attachments, lay the foundation to uncover the key mechanical and regulatory mechanisms by which kinetochores control chromosome segregation and cell division.</p>
},
	issn = {1545-9985},
	doi = {10.1038/nsmb.2358},
	author = {Gonen, Shane and Akiyoshi, Bungo and Iadanza, Matthew G and Shi, Dan and Duggan, Nicole and Biggins, Sue and Gonen, Tamir}
}
@article {3631,
	title = {Tension directly stabilizes reconstituted kinetochore-microtubule attachments.},
	journal = {Nature},
	volume = {468},
	year = {2010},
	month = {2010 Nov 25},
	pages = {576-9},
	abstract = {<p>Kinetochores are macromolecular machines that couple chromosomes to dynamic microtubule tips during cell division, thereby generating force to segregate the chromosomes. Accurate segregation depends on selective stabilization of correct {\textquoteright}bi-oriented{\textquoteright} kinetochore-microtubule attachments, which come under tension as the result of opposing forces exerted by microtubules. Tension is thought to stabilize these bi-oriented attachments indirectly, by suppressing the destabilizing activity of a kinase, Aurora B. However, a complete mechanistic understanding of the role of tension requires reconstitution of kinetochore-microtubule attachments for biochemical and biophysical analyses in vitro. Here we show that native kinetochore particles retaining the majority of kinetochore proteins can be purified from budding yeast and used to reconstitute dynamic microtubule attachments. Individual kinetochore particles maintain load-bearing associations with assembling and disassembling ends of single microtubules for \&gt;30 min, providing a close match to the persistent coupling seen in vivo between budding yeast kinetochores and single microtubules. Moreover, tension increases the lifetimes of the reconstituted attachments directly, through a catch bond-like mechanism that does not require Aurora B. On the basis of these findings, we propose that tension selectively stabilizes proper kinetochore-microtubule attachments in vivo through a combination of direct mechanical stabilization and tension-dependent phosphoregulation.</p>
},
	keywords = {Chromosomes, Fungal Proteins, Kinetochores, Microtubules, Saccharomyces cerevisiae},
	issn = {1476-4687},
	doi = {10.1038/nature09594},
	author = {Akiyoshi, Bungo and Sarangapani, Krishna K and Powers, Andrew F and Nelson, Christian R and Reichow, Steve L and Arellano-Santoyo, Hugo and Gonen, Tamir and Ranish, Jeffrey A and Asbury, Charles L and Biggins, Sue}
}
