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Model of TITAN gene functions in Arabidopsis. Nuclear division in the developing endosperm requires at least two networks of TITAN proteins. One modulates chromosome integrity through scaffold proteins known as cohesins (SMC1 and SMC3) and condensins (SMC2). Another regulates microtubule assembly through interactions between ARL2 and tubulin-folding cofactor D. Knockouts in AtSMC4, protein targets of TTN6 activity, and cellular functions of TTN4 and TTN9 remain to be identified.
