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• Drought-induced xylem embolism is a primary cause of plant mortality. Although ~70% of cycads are threatened by extinction and extant cycads diversified during a period of increasing aridification, the vulnerability of cycads to embolism spread has been overlooked. • We quantified the vulnerability to drought-induced embolism, pressure-volume curves, in situ water potentials, and a suite of xylem anatomical traits of leaf pinnae and rachises for 20 cycad species. We tested whether anatomical traits were linked to hydraulic safety in cycads. • Compared to other major vascular plant clades, cycads exhibited similar embolism resistance to angiosperms and pteridophytes but were more vulnerable to embolism than non-cycad gymnosperms. All 20 cycads had both tracheids and vessels, the proportions of which were unrelated to embolism resistance. Only vessel pit membrane fraction was positively correlated to embolism resistance, contrary to angiosperms. Water potential at turgor loss was significantly correlated to embolism resistance among cycads. • Our results show that cycads exhibit low resistance to xylem embolism and that xylem anatomical traits–particularly vessels–may influence embolism resistance together with tracheids. This study highlights the importance of understanding the mechanisms of drought resistance in evolutionarily unique and threatened lineages like the cycads. 

Flowers are critical for successful reproduction and have been a major axis of diversification among angiosperms. As the frequency and severity of droughts are increasing globally, maintaining water balance of flowers is crucial for food security and other ecosystem services that rely on flowering. Yet remarkably little is known about the hydraulic strategies of flowers. We characterized hydraulic strategies of leaves and flowers of ten species by combining anatomical observations using light and scanning electron microscopy with measurements of hydraulic physiology (minimum diffusive conductance ( g min ) and pressure-volume (PV) curves parameters). We predicted that flowers would exhibit higher g min and higher hydraulic capacitance than leaves, which would be associated with differences in intervessel pit traits because of their different hydraulic strategies. We found that, compared to leaves, flowers exhibited: 1) higher g min , which was associated with higher hydraulic capacitance ( C T ); 2) lower variation in intervessel pit traits and differences in pit membrane area and pit aperture shape; and 3) independent coordination between intervessel pit traits and other anatomical and physiological traits; 4) independent evolution of most traits in flowers and leaves, resulting in 5) large differences in the regions of multivariate trait space occupied by flowers and leaves. Furthermore, across organs intervessel pit trait variation was orthogonal to variation in other anatomical and physiological traits, suggesting that pit traits represent an independent axis of variation that have as yet been unquantified in flowers. These results suggest that flowers, employ a drought-avoidant strategy of maintaining high capacitance that compensates for their higher g min to prevent excessive declines in water potentials. This drought-avoidant strategy may have relaxed selection on intervessel pit traits and allowed them to vary independently from other anatomical and physiological traits. Furthermore, the independent evolution of floral and foliar anatomical and physiological traits highlights their modular development despite being borne from the same apical meristem. 

Abstract The leaf economics spectrum (LES) characterizes a tradeoff between building a leaf for durability versus for energy capture and gas exchange, with allocation to leaf dry mass per projected surface area (LMA) being a key trait underlying this tradeoff. However, regardless of the biomass supporting the leaf, high rates of gas exchange are typically accomplished by small, densely packed stomata on the leaf surface, which is enabled by smaller genome sizes. Here, we investigate how variation in genome size‐cell size allometry interacts with variation in biomass allocation (i.e. LMA) to influence the maximum surface conductance to CO₂and the rate of resource turnover as measured by leaf water residence time. We sampled both evergreen and deciduousRhododendron(Ericaceae) taxa from wild populations and botanical gardens, including naturally occurring putative hybrids and artificially generated hybrids. We measured genome size, anatomical traits related to cell sizes, and morphological traits related to water content and dry mass allocation. Consistent with the LES, higher LMA was associated with slower water residence times, and LMA was strongly associated with leaf thickness. Although anatomical and morphological traits varied orthogonally to each other, cell size had a pervasive impact on leaf functional anatomy: for a given leaf thickness, reducing cell size elevated the leaf surface conductance and shortened the mean water residence time. These analyses clarify how anatomical traits related to genome size‐cell size allometry can influence leaf function independently of morphological traits related to leaf longevity and durability. 

Abstract Interconduit pit membranes, which are permeable regions in the primary cell wall that connect to adjacent conduits, play a crucial role in water relations and the movement of nutrients between xylem conduits. However, how pit membrane characteristics might influence water‐carbon coupling remains poorly investigated in cycads. We examined pit characteristics, the anatomical and photosynthetic traits of 13 cycads from a common garden, to determine if pit traits and their coordination are related to water relations and carbon economy. We found that the pit traits of cycads were highly variable and that cycads exhibited a similar tradeoff between pit density and pit area as other plant lineages. Unlike other plant lineages (1) pit membranes, pit apertures, and pit shapes of cycads were not coordinated as in angiosperms; (2) cycads exhibited larger pit membrane areas but lower pit densities relative to ferns and angiosperms, but smaller and similar pit membrane densities to non‐cycad gymnosperms; (3) cycad pit membrane areas and densities were partially coordinated with anatomical traits, with hydraulic supply of the rachis positively coordinated with photosynthesis, whereas pit aperture areas and fractions were negatively coordinated with photosynthetic traits; (4) cycad pit traits reflected adaptation to wetter habitats for Cycadaceae and drier habitats for Zamiaceae. The large variation in pit traits, the unique pit membrane size and density, and the partial coordination of pit traits with anatomical and physiological traits of the rachis and pinna among cycads may have facilitated their dominance in a variety of ecosystems from the Mesozoic to modern times.