The discovery of visible fluorescence in the plant pigments betalains revealed the existence of fluorescent patterns in flowers of plants of the order Caryophyllales, where betalains substitute anthocyanins. The serendipitous initial discovery led to a systemized characterization of the role of different substructures on the photophysical phenomenon. Strong fluorescence is general to all members of the family of betaxanthins linked to the structural property that the betalamic acid moiety is connected to an amine group. This property has led to bioinspired tailor-made probes and to the development of novel biotechnological applications in screening techniques or microscopy labeling. Here, we comprehensively review the photophysics, photochemistry, and photobiology of betalain fluorescence and describe all current applications.
BetalainsBetalains are nitrogenous plant pigments that are characteristic for plants belonging to the order Caryophyllales. These pigments are divided into the yellow betaxanthins and the violet betacyanins [1]. The presence of both types of pigments is required for the orange and red colors that coexist in nature with the pure yellow and violet colors. Betalains substitute anthocyanins and play their roles in the colored tissues of most plant families of the Caryophyllales. Both families of pigments are watersoluble and mutually exclusive [2,3]. Numerous studies were performed on the color properties of betalains since they were described as a novel family of pigments, different to 'nitrogenous anthocyanins' [4,5]. However, it was not until 2005 that their natural fluorescence was discovered [6]. In addition, it was demonstrated that under physiological conditions, light emission was exhibited in the plant tissues that contain them, including flowers [7,8].Among the Caryophyllales plants, red beet roots (Beta vulgaris) and the fruits of cacti belonging to the genus Opuntia are the best known sources of betalains, with betanin and indicaxanthin being, respectively, their main pigments [9,10]. Current research has described the betalain content of novel sources, such as the tubers from Ullucus tuberosus [11] or the betalain-containing berries of Rivina humilis [12]. Also, the multiple shades of quinoa grains (Chenopodium quinoa) have recently been reported to be based on betacyanins and betaxanthins [13]. In addition, betalains have in recent years been shown to have promising bioactive properties. Early investigations revealed a strong free radical scavenging capacity of betalains purified from beet root [14]. Subsequent research revealed the existence of an intrinsic activity, present in all betalains, that is modulated by structural factors [15,16]. Furthermore, studies with different human cancer cell lines have demonstrated the potential of betalains in the chemoprevention of cancer [17,18]. In vivo experiments have shown that very low concentrations of dietary pigments inhibit the formation of tumors in mice [19,20] and extend the lifespan of the model animal Caenorhabditis elegans [21...