RESUMO
The onset of stomatal closure reduces transpiration during drought. In seed plants, drought causes declines in plant water status which increases leaf endogenous abscisic acid (ABA) levels required for stomatal closure. There are multiple possible points of increased belowground resistance in the soil-plant atmospheric continuum that could decrease leaf water potential enough to trigger ABA production and the subsequent decreases in transpiration. We investigate the dynamic patterns of leaf ABA levels, plant hydraulic conductance and the point of failure in the soil-plant conductance in the highly embolism-resistant species Callitris tuberculata using continuous dendrometer measurements of leaf water potential during drought. We show that decreases in transpiration and ABA biosynthesis begin before any permanent decreases in predawn water potential, collapse in soil-plant hydraulic pathway and xylem embolism spread. We find that a dynamic but recoverable increases in hydraulic resistance in the soil in close proximity to the roots is the most likely driver of declines in midday leaf water potential needed for ABA biosynthesis and the onset of decreases in transpiration.
RESUMO
The phytohormone abscisic acid (ABA) is synthesised by plants during drought to close stomata and regulate desiccation tolerance pathways. Conifers and some angiosperms with embolism-resistant xylem show a peaking-type (p-type) response in ABA levels, in which ABA levels increase early in drought then decrease as drought progresses, declining to pre-stressed levels. The mechanism behind this dynamic remains unknown. Here, we sought to characterise the mechanism driving p-type ABA dynamics in the conifer Callitris rhomboidea and the highly drought-resistant angiosperm Umbellularia californica. We measured leaf water potentials (Ψl ), stomatal conductance, ABA, conjugates and phaseic acid (PA) levels in potted plants during a prolonged but non-fatal drought. Both species displayed a p-type ABA dynamic during prolonged drought. In branches collected before and after the peak in endogenous ABA levels in planta, that were rehydrated overnight and then bench dried, ABA biosynthesis was deactivated beyond leaf turgor loss point. Considerable conversion of ABA to conjugates was found to occur during drought, but not catabolism to PA. The mechanism driving the decline in ABA levels in p-type species may be conserved across embolism-resistant seed plants and is mediated by sustained conjugation of ABA and the deactivation of ABA accumulation as Ψl becomes more negative than turgor loss.
Assuntos
Embolia , Magnoliopsida , Traqueófitas , Estômatos de Plantas/fisiologia , Secas , Folhas de Planta/metabolismo , Ácido Abscísico/metabolismo , Água/metabolismo , Magnoliopsida/fisiologiaRESUMO
The stems of some herbaceous species can undergo basal secondary growth, leading to a continuum in the degree of woodiness along the stem. Whether the formation of secondary growth in the stem base results in differences in embolism resistance between the base and the upper portions of stems is unknown. We assessed the embolism resistance of leaves and the basal and upper portions of stems simultaneously within the same individuals of two divergent herbaceous species that undergo secondary growth in the mature stem bases. The species were Solanum lycopersicum (tomato) and Senecio minimus (fireweed). Basal stem in mature plants of both species displayed advanced secondary growth and greater resistance to embolism than the upper stem. This also resulted in significant vulnerability segmentation between the basal stem and the leaves in both species. Greater embolism resistance in the woodier stem base was found alongside decreases in the pith-to-xylem ratio, increases in the proportion of secondary xylem, and increases in lignin content. We show that there can be considerable variation in embolism resistance across the stem in herbs and that this variation is linked to the degree of secondary growth present. A gradient in embolism resistance across the stem in herbaceous plants could be an adaptation to ensure reproduction or basal resprouting during episodes of drought late in the lifecycle.
Assuntos
Folhas de Planta , Caules de Planta , Caules de Planta/crescimento & desenvolvimento , Caules de Planta/fisiologia , Folhas de Planta/crescimento & desenvolvimento , Folhas de Planta/fisiologia , Xilema/fisiologia , Xilema/crescimento & desenvolvimento , Solanum lycopersicum/crescimento & desenvolvimento , Solanum lycopersicum/fisiologia , Lignina/metabolismo , Combretaceae/fisiologia , Combretaceae/crescimento & desenvolvimentoRESUMO
Maintaining water transport in the xylem is critical for vascular plants to grow and survive. The drought-induced accumulation of embolism, when gas enters xylem conduits, causes declines in hydraulic conductance (K ) and is ultimately lethal. Several methods can be used to estimate the degree of embolism in xylem, from measuring K in tissues to directly visualising embolism in conduits. One method allowing a direct quantification of embolised xylem area is the optical vulnerability (OV) technique. This method has been used across different organs and has a high spatial and temporal resolution. Here, we review studies using the OV technique, discuss the main advantages and disadvantages of this method, and summarise key advances arising from its use. Vulnerability curves generated by the OV method are regularly comparable to other methods, including the centrifuge and X-ray microtomography. A major advantage of the OV technique over other methods is that it can be simultaneously used to determine in situ embolism formation in leaves, stems and roots, in species spanning the phylogeny of land plants. The OV method has been used to experimentally investigate the spreading of embolism through xylem networks, associate embolism with downstream tissue death, and observe embolism formation in the field.