Relative expression of the two FAZYs, 1-SST, 1-FFT along with their corresponding metabolite (fructans, sucrose, glucose) were assessed in both artichoke and wild cardoon sampled at stage 5 (inflorescence emergence and development of the head) (Fig. 1). The inulin-type fructans occurred in the rhizome of both taxa with a slight lower level in the wild cardoon rhizome (64,72 ± 7,71 g/100g) respect with artichoke one (80,03 ± 7,03 g/100g); conversely a much higher quantity was detected in the heads of the wild cardoon (72,53 ± 12,03 g/100g) compared with artichoke ones (34,03 ± 9,85 g/100g) (Fig. 2A). The WSC, considered as mono (glucose, fructose) and disaccharides (sucrose) fractions, showed a significative higher content in wild cardoon heads (20.70 ± 8.03g/100g) compared with artichoke ones (14.49 ± 3.83g/100g) (Fig. 2B); also, in the wild cardoon rhizome a higher WSC level (25.64 ± 6.05g/100g) vs artichoke (12.95 ± 4.67g/100g) was detected (Fig. 2B).
The transcript levels of 1-SST and 1-FFT showed generally a higher expression in heads than rhizomes with significant higher level in wild cardoon head respect with globe artichoke ones (Fig. 3); conversely the downregulation of the expression of both enzymes observed in the rhizomes was not statistically significant between the two taxa (Fig. 3).
The high activity of 1-SST and 1-FFT associated with a higher concentration of fructans in the wild cardoon head, supports the hypothesis that fructans are produced by the active action of 1-SST /1-FFT enzyme; whereas in the rhizome, where the expression of 1-SST /1-FFT enzyme are low and at a basal level, fructans are probably the residual of the storage material that in these organs occurred during the resting season. Indeed, in some plants accumulation of fructans is observed under severe drought and cold stresses, since they are resistant to crystallization at low temperatures (Krasensky and Jonak 2012); additionally, acting as a source of reserve carbohydrate, they can get utilized during the recovery phase after a stress (Konstantinova et al. 2002). Moreover, the high availability of WSC in the wild cardoon, depending mainly on the release of glucose and sucrose, can be explained by the increase in 1-SST enzyme activity (Cairns 1993); in fact a good correlation between concentrations of free glucose (one of the products of 1-SST) with expression level of 1-SST gene was approved by Van Den Ende et al. (1996). Although the exact regulatory pathways that control fructans metabolism are not fully known and they are a subject of ongoing research in several laboratories, it reasonable expected that the pattern of storage polysaccharide accumulation is a dynamic process intimately interconnected with those controlling storage organs differentiation (in this case the rhizome) and plant growth. In fact, energetic substances accumulate in rhizome ensure the supply of compounds when new growth cycle starts after the summer quiescence (Raccuia and Melilli 2010; Gominho et al. 2018). However, it can be expected that genes for fructans/inulin biosynthetic and breakdown enzymes will be also regulated developmentally by sugars, wounding, hormones and stress factors such as, for example, drought or low temperature (Joudi et al. 2012; Livingston et al. 2009).
It is interesting to note that at the stage of head development, the wild heads cardoon is much richer in fructans than artichoke ones; we can assume that this carbohydrate reserve in the form of fructans could play an important role in stressful conditions. Probably this wild genotype retains some genetic traits conferring it a greater adaptability to marginal area and coping with stress much more than cultivated artichoke. So, the great fructans levels observed during the head development, confer to the wild cardoon a good potential for health promoting dietary aliments and also it can be a good alternative to the common and already greatly commercialized artichoke; overall the exploitation of wild cardoon for the application in both food and industrial sectors could be more promising as suitable crop of marginal and harsh Mediterranean area. A further interesting note is that the expression pathway of 1-SST /1-FFT keys enzymes may be an early marker of fructans biosynthesis and therefore a good label of production of these interesting metabolites also at the industrial level; nevertheless, this two FAZYs may be a valuable fast tool to identify suitable cultivars and/or more promising genotypes in breeding or conservation program of industrial artichoke.