During mitosis, chromosomes become attached to the structure known as the mitotic spindle. In the late 1800s, Theodor Boveri created the earliest detailed drawings of the spindle based on his observations of cell division in early Ascaris embryos. Boveri's drawings, which are amazingly accurate, show chromosomes attached to a bipolar network of fibers.. We now know that centrioles duplicate during S phase, although many details of this duplication process are still under investigation. The composition of the spindle fibers remained unknown until the 1960s, when tubulin was discovered and techniques were developed for visualizing spindles using electron microscopes. The length of these kinetochore-attached microtubules then decreases during mitosis, pulling sister chromatids to opposite poles of the spindle.
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