{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\nTTK (also known as Mps1) is a dual‐specificity protein kinase that plays a central role in ensuring accurate chromosome segregation during mitosis. It orchestrates the spindle assembly checkpoint (SAC) by localizing to kinetochores and recruiting key checkpoint proteins—including Mad1, Mad2, Bub1, BubR1, and others—to monitor kinetochore–microtubule attachment and correct improper attachments. TTK’s kinase activity is required for error correction as it works downstream of Aurora B and collaborates with components of the Ndc80 complex to sense attachment status. Moreover, TTK fosters timely checkpoint engagement and release, thus regulating both the onset of anaphase and the fidelity of centrosome duplication."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "1", "end_ref": "13"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nIn addition to its core checkpoint function, TTK activity is tightly regulated by autophosphorylation and by phosphorylation of downstream substrates. It interacts with several other kinases and regulatory proteins—including Aurora B, CHK2, and components of the TGF‐β/Smad pathway—to modulate kinetochore dynamics and checkpoint silencing. Its own activation via trans autophosphorylation (for example on residues within the activation loop) is crucial for both its catalytic efficiency and its timely release from kinetochores upon correct biorientation. Such post‐translational modifications and protein–protein interactions, often mediated by discrete structural modules within TTK, ensure that mitotic progression is exquisitely responsive to microtubule attachment status."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "14", "end_ref": "32"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nTTK is also emerging as an attractive target in oncology. Elevated expression of TTK has been observed in multiple cancer types—including triple‐negative breast, colon, and hepatocellular carcinomas—and correlates with genomic instability, poor prognosis, and therapeutic resistance. Inhibition or depletion of TTK not only disrupts checkpoint signaling but also impairs cell proliferation, sensitizes tumor cells to chemotherapy or radiation, and can drive mitotic catastrophe. 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