Akusala-mula, translated as 'roots of unwholesomeness,' refers to the three mental states that are considered the primary causes of unwholesome actions leading to suffering. These roots include greed (lobha), hatred (dosa), and delusion (moha). Understanding akusala-mula is essential in grasping how these negative mental states contribute to the cycle of suffering and how they can be transformed into their wholesome counterparts for spiritual development.