As we face the millenium, a new and more integrated view of the cytoskeleton has begun to emerge. While each cytoskeleton retains its identity, the three networks rely heavily upon one another, employing connections for cytoskeletal stability, intracellular transport and trafficking, cytokinesis, cell polarity, and tissue morphogenesis. Key genetic evidence underscores the importance of connector proteins to cytoarchitecture and structural integrity as well as many dynamic cellular processes including migration. The emergence of plakins as a vital group of linker proteins that bridge IFs, MTs, and/or MF networks has paved the entrance to a remarkable interstate of novel functional interactions among the structural elements within the cytoplasm. The discovery of actin-MT connections as well as plakins in lower eukaryotes that lack IFs, suggest that the foundations for these fascinating new cellular pathways have ancient origins.