Injury to the xylem and vascular cambium is proposed to explain mortality following low severity fires. These tissues have been assessed independently, but the relative significance of the xylem and cambium is still uncertain. The goal of this study is to evaluate the xylem dysfunction hypothesis and cambium necrosis hypothesis simultaneously. The hot dry conditions of a low severity fire were simulated in a drying oven, exposing Sequoia sempervirens (Lamb. ex D. Don) shoots to 70 and 100 °C for 6–60 min. Cambial viability was measured with Neutral Red stain and water transport capacity was assessed by calculating the loss of hydraulic conductivity. Vulnerability curves were also constructed to determine susceptibility to drought-induced embolism following heat exposure. The vascular cambium died completely at 100 °C after only 6 min of heat exposure, while cells remained viable at 70 °C temperatures for up to 15 min. Sixty minutes of exposure to 70 °C reduced stem hydraulic conductivity by 40%, while 45 min at 100 °C caused complete loss of conductivity. The heat treatments dropped hydraulic conductivity irrecoverably but did not significantly impact post-fire vulnerability to embolism. Overall, the damaging effects of high temperature occurred more rapidly in the vascular cambium than xylem following heat exposure. Importantly, the xylem remained functional until the most extreme treatments, long after the vascular cambium had died. Our results suggest that the viability of the vascular cambium may be more critical to post-fire survival than xylem function in S. sempervirens. Given the complexity of fire, we recommend ground-truthing the cambial and xylem post-fire response on a diverse range of species.
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Abstract Background and Aims The success of invasive plants can be attributed to many traits including the ability to adapt to variable environmental conditions. Whether by adaptation, acclimation or phenotypic plasticity, these plants often increase their resource-use efficiency and, consequently, their fitness. The goal of this study was to examine the hydraulic and eco-physiological attributes of sun and shade populations of Pteridium aquilinum, a weedy fern, to determine whether the presence of vessels and other hydraulic attributes affects its success under a variety of light conditions. Methods Hydraulic traits such as cavitation resistance, hydraulic conductivity, photosynthesis and water potential at turgor loss point were measured on fronds from sun and shade populations. Anatomical and structural traits such as conduit diameter and length, stomatal density and vein density were also recorded. Diurnal measures of leaf water potential and stomatal conductance complement these data. Key Results Gas exchange was nearly double in the sun plants, as was water-use efficiency, leaf-specific conductivity, and stomatal and vein density. This was largely achieved by a decrease in leaf area, coupled with higher xylem content. There was no significant difference in petiole cavitation resistance between the sun and shade leaves, nor in xylem-specific conductivity. Hydraulic conduit diameters were nearly equivalent in the two leaf types. Conclusions Shifts in leaf area and xylem content allow P. aquilinum to occupy habitats with full sun, and to adjust its physiology accordingly. High rates of photosynthesis explain in part the success of this fern in disturbed habitats, although no change was observed in intrinsic xylem qualities such as cavitation resistance or conduit length. This suggests that P. aquilinum is constrained by its fundamental body plan, in contrast to seed plants, which show greater capacity for hydraulic adjustment.more » « less
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Summary The Cretaceous–Cenozoic expansion of tropical forests created canopy space that was subsequently occupied by diverse epiphytic communities including Eupolypod ferns. Eupolypods proliferated in this more stressful niche, where lower competition enabled the adaptive radiation of thousands of species. Here, we examine whether xylem traits helped shape the Cenozoic radiation of Eupolypod ferns.
We characterized the petiole xylem anatomy of 39 species belonging to the Eupolypod I and Eupolypod II clades occupying the epiphytic, hemiepiphytic, and terrestrial niche, and we assessed vulnerability to embolism in a subset of species.
The transition to the canopy was associated with reduced xylem content and smaller tracheid diameters, but no differences were found in species vulnerability to embolism and pit membrane thickness. Phylogenetic analyses support selection for traits associated with reduced water transport in Eupolypod 1 species.
We posit that in Eupolypod epiphytes, selection favored water retention via thicker leaves and lower stomatal density over higher rates of water transport. Consequently, lower leaf water loss was coupled with smaller quantities of xylem and narrower tracheid diameters. Traits associated with water conservation were evident in terrestrial Eupolypod 1 ferns and may have predisposed this clade toward radiation in the canopy.
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Summary Different methods of measuring cavitation resistance in fern petioles lead to variable results, particularly with respect to the P50metric. We hypothesised that the fern dictyostele structure affects air entry into the xylem, and therefore impacts the shape of the vulnerability curve.
Our study examined this variation by comparing vulnerability curves constructed on petioles collected from evergreen and deciduous ferns in the field, with curves generated using the standard centrifuge, air‐injection and bench‐top dehydration methods. Additional experiments complemented the vulnerability curves to better understand how anatomy shapes estimates of cavitation resistance.
Centrifugation and radial air injection generated acceptable vulnerability curves for the deciduous species, but overestimated drought resistance in the two evergreen ferns. In these hardy plants, axial air injection and bench‐top dehydration produced results that most closely aligned with observations in nature. Additional experiments revealed that the dictyostele anatomy impedes air entry into the xylem during spinning and radial air injection.
Each method produced acceptable vulnerability curves, depending on the species being tested. Therefore, we stress the importance of validating the curves with
in situ measures of water potential and, if possible, hydraulic data to generate realistic results with any of the methods currently available. -
Abstract Desiccation-tolerant (DT) organisms can lose nearly all their water without dying. Desiccation tolerance allows organisms to survive in a nearly completely dehydrated, dormant state. At the cellular level, sugars and proteins stabilize cellular components and protect them from oxidative damage. However, there are few studies of the dynamics and drivers of whole-plant recovery in vascular DT plants. In vascular DT plants, whole-plant desiccation recovery (resurrection) depends not only on cellular rehydration, but also on the recovery of organs with unequal access to water. In this study, in situ natural and artificial irrigation experiments revealed the dynamics of desiccation recovery in two DT fern species. Organ-specific irrigation experiments revealed that the entire plant resurrected when water was supplied to roots, but leaf hydration alone (foliar water uptake) was insufficient to rehydrate the stele and roots. In both species, pressure applied to petioles of excised desiccated fronds resurrected distal leaf tissue, while capillarity alone was insufficient to resurrect distal pinnules. Upon rehydration, sucrose levels in the rhizome and stele dropped dramatically as starch levels rose, consistent with the role of accumulated sucrose as a desiccation protectant. These findings provide insight into traits that facilitate desiccation recovery in dryland ferns associated with chaparral vegetation of southern California.more » « less
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Xylem form and function under extreme nutrient limitation: an example from California's pygmy forestmore » « less
Summary Xylem anatomy and function have large implications for plant growth as well as survival during drought, but the impact of nutrient limitation on xylem is not fully understood. This study examines the pygmy forest in California, a plant community that experiences negligible water stress but is severely stunted by low‐nutrient and acidic soil, to investigate how nutrient limitation affects xylem function.
Thirteen key anatomical and hydraulic traits of stems of four species were compared between pygmy forest plants and nearby conspecifics growing on richer soil.
Resistance to cavitation (P50), a critical trait for predicting survival during drought, had highly species‐specific responses: in one species, pygmy plants had a 26% decrease in cavitation resistance compared to higher‐nutrient conspecifics, while in another species, pygmy plants had a 56% increase in cavitation resistance. Other traits responded to nutrient limitation more consistently: pygmy plants had smaller xylem conduits and higher leaf‐specific conductivity (
K L) than conspecific controls.Edaphic stress, even in the absence of water stress, altered xylem structure and thus had substantial impacts on water transport. Importantly, nutrient limitation shifted cavitation resistance in a species‐specific and unpredictable manner; this finding has implications for the assessment of cavitation resistance in other natural systems.
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Abstract Opportunistic diversification has allowed ferns to radiate into epiphytic niches in angiosperm dominated landscapes. However, our understanding of how ecophysiological function allowed establishment in the canopy and the potential transitionary role of the hemi‐epiphytic life form remain unclear. Here, we surveyed 39 fern species in Costa Rican tropical forests to explore epiphytic trait divergence in a phylogenetic context. We examined leaf responses to water deficits in terrestrial, hemi‐epiphytic and epiphytic ferns and related these findings to functional traits that regulate leaf water status. Epiphytic ferns had reduced xylem area (−63%), shorter stipe lengths (−56%), thicker laminae (+41%) and reduced stomatal density (−46%) compared to terrestrial ferns. Epiphytic ferns exhibited similar turgor loss points, higher osmotic potential at saturation and lower tissue capacitance after turgor loss than terrestrial ferns. Overall, hemi‐epiphytic ferns exhibited traits that share characteristics of both terrestrial and epiphytic species. Our findings clearly demonstrate the prevalence of water conservatism in both epiphytic and hemi‐epiphytic ferns, via selection for anatomical and structural traits that avoid leaf water stress. Even with likely evolutionarily constrained physiological function, adaptations for drought avoidance have allowed epiphytic ferns to successfully endure the stresses of the canopy habitat.
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Summary Desiccation‐tolerant (DT) plants can dry past −100 MPa and subsequently recover function upon rehydration. Vascular DT plants face the unique challenges of desiccating and rehydrating complex tissues without causing structural damage. However, these dynamics have not been studied in intact DT plants.
We used high resolution micro‐computed tomography (microCT), light microscopy, and fluorescence microscopy to characterize the dynamics of tissue desiccation and rehydration in petioles (stipes) of intact DT ferns.
During desiccation, xylem conduits in stipes embolized before cellular dehydration of living tissues within the vascular cylinder. During resurrection, the chlorenchyma and phloem within the stipe vascular cylinder rehydrated before xylem refilling. We identified unique stipe traits that may facilitate desiccation and resurrection of the vascular system, including xylem conduits containing pectin (which may confer flexibility and wettability); chloroplasts within the vascular cylinder; and an endodermal layer impregnated with hydrophobic substances that impede apoplastic leakage while facilitating the upward flow of water within the vascular cylinder.
Resurrection ferns are a novel system for studying extreme dehydration recovery and embolism repair in the petioles of intact plants. The unique anatomical traits identified here may contribute to the spatial and temporal dynamics of water movement observed during desiccation and resurrection.
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Summary Sex expression of homosporous ferns is controlled by multiple factors, one being the antheridiogen system. Antheridiogens are pheromones released by sexually mature female fern gametophytes, turning nearby asexual gametophytes precociously male. Nevertheless, not all species respond. It is still unknown how many fern species use antheridiogens, how the antheridiogen system evolved, and whether it is affected by polyploidy and/or apomixis.
We tested the response of 68 fern species to antheridiogens in cultivation. These results were combined with a comprehensive review of literature to form the largest dataset of antheridiogen interactions to date. Analyzed species also were coded as apomictic or sexual and diploid or polyploid.
Our final dataset contains a total of 498 interactions involving 208 species (
c . 2% of all ferns). About 65% of studied species respond to antheridiogen. Multiple antheridiogen types were delimited and their evolution is discussed. Antheridiogen responsiveness was not significantly affected by apomixis or polyploidy.Antheridiogens are widely used by ferns to direct sex expression. The antheridiogen system likely evolved multiple times and provides homosporous ferns with the benefits often associated with heterospory, such as increased rates of outcrossing. Despite expectations, antheridiogens may be beneficial to polyploids and apomicts.
