“…Elucidation of these core enzymatic steps has enabled the engineering of the betalain biosynthetic pathway in a wide range of heterologous hosts, including microbes such as Saccharomyces cerevisiae [38]; plant model organisms like Arabidopsis thaliana, Nicotiana tabacum, and Petunia hybrida [31,39,40]; and a diversity of crops such as Oryza sativa (rice), Solanum lycopersicum (tomato), Solanum tuberosum (potato), and Solanum melongena (aubergine) [40,41]. Betalains have been used as biosensors in a number of heterologous contexts to report increased production of metabolites, such as tyrosine, dopamine, and ʟ-DOPA in Escherichia coli and Nicotiana benthamiana [23,24,26], to measure metabolic flux between competing pathways in the synthesis of benzylisoquinoline alkaloids (BIAs) in S. cerevisiae [22] and for the detection of copper by heavy metal-resistant bacteria in bioremediation processes [25]. In plants, specific promoters have been successfully used to target betalain production in specific tissues such as fruits and seed endosperm [40][41][42].…”