From cells to stems: the effects of primary vascular construction on drought-induced embolism in fern rhizomes

While a considerable amount of data exists on the link between xylem construction and hydraulic function, few studies have focused on resistance to drought-induced embolism of primary vasculature in herbaceous plants. Ferns rely entirely on primary xylem and display a remarkable diversity of vascular construction in their rhizomes, making them an ideal group in which to examine hydraulic structure-function relationships. New optical methods allowed us to measure vulnerability to embolism in rhizomes, which are notoriously difficult to work with. We investigated five fern species based on their diverse xylem traits at the cellular, histological, and architectural levels. To link below- and above-ground hydraulics, we then measured leaf-stem vulnerability segmentation. Overall, rhizome vulnerability to embolism was correlated most strongly with cellular but not histological or architectural traits. Interestingly, at P6-12, species with increased architectural dissection were actually more vulnerable to embolism, suggesting different hydraulic dynamics at low compared to high percent embolism. Importantly, leaves fully embolize before stems reach P-88, suggesting strong vulnerability segmentation. This is the first study to explore the functional implications of primary vascular construction in fern rhizomes and leaf-stem vulnerability segmentation. Strong segmentation suggests that leaves protect perennial rhizomes against severe drought stress and hydraulically induced mortality.