Contractile roots in succulent monocots: convergence, divergence and adaptation to limited rainfall.

Contractile roots (CRs) that pull shoots further down in the soil are a possible example of convergent evolution in two monocot families, the Agavaceae and the Asphodelaceae. The association between CRs, water uptake and habitat aridity was investigated for agaves, yuccas and aloes by assessing the occurrence of CRs and the amount of root contraction for glasshouse-grown plants with respect to mean annual rainfall of their native habitats. Structural features of CRs as well as root hydraulic conductance were compared with those of non-contractile roots (NCRs). CRs occurred in 55% of the 73 species examined, including 64% of the agaves and 85% of the yuccas, but in none of the aloes despite the occurrence of CRs in related genera. The phylogenetic distribution of CRs was consistent with multiple acquisitions or losses of the trait. The amount of root contraction showed a highly significant negative relationship with mean annual rainfall, although other environmental factors may also be important. Radial hydraulic conductance of the basal (contractile) zone exceeded that of the midroot zone for CRs; for NCRs, the opposite was true. Thus, CRs in the species examined may provide a mechanism for greater water uptake near the soil surface in regions with limited rainfall.

Root contraction helps protect the "living rock" cactus Ariocarpus fissuratus from lethal high temperatures when growing in rocky soil.

PREMISE OF THE STUDY: We investigated how the "living rock" cactus Ariocarpus fissuratus, like other low-growing desert plants, can endure potentially lethal high temperatures at the soil surface. Specifically, we examined how shoot descent by root contraction in the presence or absence of soil rocks influences shoot temperatures and transpiration. • METHODS: Root contraction was identified by measuring shoot descent and anatomical analysis. Temperatures and transpiration were measured for plants at two heights in sandy and rocky soil, and temperature tolerances were determined by vital staining. • KEY RESULTS: Plants embedded in rocky soil survived an extreme heat episode, unlike plants in sandy soil, though rocks did not moderate low temperatures. Root contraction occurred regardless of season and soil moisture. Xylem conduits (wide-band tracheids) formed a compressible lattice that decreased root length as rays enlarged the root base radially. Plant position in the soil did not affect transpiration. • CONCLUSIONS: Contractile roots pulled plants of A. fissuratus into the soil at rates of 6-30 mm yr(-1). Maintaining shoots level with the soil surface kept plant temperatures below the high lethal temperature and improved survivorship in soil shaded by surface rocks.