(A) In fungal model systems, cortically anchored dynein-dynactin complexes move the nucleus toward the mother-bud neck (the future site of cytokinesis) in budding yeast (left), or in the direction of the growing hyphal tip to ensure proper nuclear distribution in Aspergillus nidulans (right). Dynein-dynactin complexes are recruited and anchored to the cell cortex in budding yeast and A. nidulans by Num1 and ApsA, respectively, (Fischer and Timberlake, 1995; Heil-Chapdelaine et al., 2000; Veith et al., 2005), which exhibit significant degrees of similarity within their C-terminal pleckstrin homology domains (for membrane association; 50% identity/64% similarity) and their N-terminal coiled-coil-containing dynein-dynactin interacting regions (24% identity/41% similarity; note this has not been experimentally determined in A. nidulans) (Tang et al., 2012). (B) In the developing brain, LIS1 and dynein are critical for the three distinct illustrated processes: (Box 1) During interkinetic nuclear migration (INM), the nucleus is transported to the ventricular surface in highly elongated radial glial precursors (RGPs), where mitosis takes place. Dynein-dynactin complexes anchored to the nuclear envelope – through interactions between dynein adaptors (e.g., BicD2, in yellow) and nuclear pore complex components (e.g., RanBP2, in red; nuclear pore complex, or NPC, in orange) – transport the nucleus toward the minus ends of microtubules (in red), which are situated toward the ventricular surface (‘-' and ‘+' depict minus and plus ends of microtubules, respectively). (Box 2) The mitotic spindles (in red, with chromosomes shown in blue) in RGPs are actively rotated by cortical dynein-dynactin complexes. The orientation of the spindle dictates whether a dividing cell will generate two RGPs (horizontal orientation, left), or one RGP and a differentiated cell type (e.g., a neuronal precursor, or NP cell; vertical, right). (Box 3) Subsequent to their generation from RGPs, NPs ascend toward the pial surface of the developing brain, where the cortical plate resides. Evidence suggests that dynein is enriched in a dilation, or ‘swelling’ within the leading process. It has been proposed that dynein anchored at the cell cortex (by an unknown cortical receptor, in yellow) in this region pulls on astral microtubules emanating from the centrosome, thus moving the nucleus toward the leading process (see text) (Tsai et al., 2007). The manner by which the centrosome is linked to the nucleus is not entirely clear, but may rely on dynein and microtubules (Vallee et al., 2009).