Auxin and cytokinin are key hormonal orchestrators of root system architecture and its developmental plasticity. Our research has identified several convergence points and signaling pathways that integrate auxin and cytokinin inputs to coordinate root organ development. Interestingly, some of the recently identified molecular components go beyond classical auxin–cytokinin cross-talk, establishing functional connections with other regulatory pathways. These include mechanisms mediating the perception of environmental cues, such as abiotic stress and nitrate availability, as well as the regulation of the microtubule cytoskeleton and subcellular trafficking.  Insights into how auxin and cytokinin signaling are embedded within complex molecular networks that govern plant growth and enable adaptive responses to fluctuating environmental conditions will be discussed.