Conduit position and connectivity affect the likelihood of xylem embolism during natural drought in evergreen woodland species.

BACKGROUND AND AIMS: Hydraulic failure is considered a main cause of drought-induced forest mortality. Yet, we have a limited understanding of how the varying intensities and long time scales of natural droughts induce and propagate embolism within the xylem. METHODS: X-ray computed tomography (microCT) images were obtained from different aged branch xylem to study the number, size and spatial distribution of in situ embolized conduits among three dominant tree species growing in a woodland community. KEY RESULTS: Among the three studied tree species, those with a higher xylem vulnerability to embolism (higher water potential at 50 % loss of hydraulic conductance; P50) were more embolized than species with lower P50. Within individual stems, the probability of embolism was independent of conduit diameter but associated with conduit position. Rather than the occurrence of random or radial embolism, we observed circumferential clustering of high and low embolism density, suggesting that embolism spreads preferentially among conduits of the same age. Older xylem also appeared more likely to accumulate embolisms than young xylem, but there was no pattern suggesting that branch tips were more vulnerable to cavitation than basal regions. CONCLUSIONS: The spatial analysis of embolism occurrence in field-grown trees suggests that embolism under natural drought probably propagates by air spreading from embolized into neighbouring conduits in a circumferential pattern. This pattern offers the possibility to understand the temporal aspects of embolism occurrence by examining stem cross-sections.

Herb and conifer roots show similar high sensitivity to water deficit.

Root systems play a major role in supplying the canopy with water, enabling photosynthesis and growth. Yet, much of the dynamic response of root hydraulics and its influence on gas exchange during soil drying and recovery remains uncertain. We examined the decline and recovery of the whole root hydraulic conductance (Kr) and canopy diffusive conductance (gc) during exposure to moderate water stress in two species with contrasting root systems: Tanacetum cinerariifolium (herbaceous Asteraceae) and Callitris rhomboidea (woody conifer). Optical dendrometers were used to record stem water potential at high temporal resolution and enabled non-invasive measurements of Kr calculated from the rapid relaxation kinetics of water potential in hydrating roots. We observed parallel declines in Kr and gc to <20% of unstressed levels during the early stages of water stress in both species. The recovery of Kr after rewatering differed between species. T. cinerariifolium recovered quickly, with 60% of Kr recovered within 2 h, while C. rhomboidea was much slower to return to its original Kr. Recovery of gc followed a similar trend to Kr in both species, with C. rhomboidea slower to recover. Our findings suggest that the pronounced sensitivity of Kr to drought is a common feature among different plant species, but recovery may vary depending on root type and water stress severity. Kr dynamics are proposed to modulate gc response during and following drought.

Previous drought exposure leads to greater drought resistance in eucalypts through changes in morphology rather than physiology.

Over their lifetime, trees are repeatedly exposed to droughts. It is therefore important to understand whether repeated drought exposure makes trees more or less drought tolerant. Here, we investigated the effect of repeated droughts on functional trait expression and tree function in Eucalyptus obliqua. Further, we tested whether previous drought exposure enabled trees to avoid leaf death for longer under a subsequent severe drought. Trees were subjected for 1 year to 2 drought-rewatering cycles (drought treatment) or well-watered conditions, before imposing a severe drought. Trees in the drought treatment reduced their overall leaf area and biomass, whereas leaf-level anatomical, morphological and physiological traits remained mostly unaffected. There were no differences in water potential at the turgor loss point, leaf xylem vulnerability to embolism, leaf size, maximum xylem vessel diameter or cell wall thickness between treatments after the conditioning period. When exposed to a subsequent severe drought, trees previously exposed to drought were more drought tolerant due to a lower water potential at leaf death and tree-level morphological rather than physiological adjustments. Trees previously exposed to drought were smaller and used less water, which delayed leaf death for 39 days compared with 22 days for the well-watered trees. Our study indicates that previous drought exposure can facilitate tree-level morphological adjustment, which potentially enhances survival of E. obliqua trees during subsequent drought events.

Phenotypic plasticity and genetic adaptation of functional traits influences intra-specific variation in hydraulic efficiency and safety.

Understanding which hydraulic traits are under genetic control and/or are phenotypically plastic is essential in understanding how tree species will respond to rapid shifts in climate. We quantified hydraulic traits in Eucalyptus obliqua L'Her. across a precipitation gradient in the field to describe (i) trait variation in relation to long-term climate and (ii) the short-term (seasonal) ability of traits to adjust (i.e., phenotypic plasticity). Seedlings from each field population were raised under controlled conditions to assess (iii) which traits are under strong genetic control. In the field, drier populations had smaller leaves with anatomically thicker xylem vessel walls, a lower leaf hydraulic vulnerability and a lower water potential at turgor loss point, which likely confers higher hydraulic safety. Traits such as the water potential at turgor loss point and ratio of sapwood to leaf area (Huber value) showed significant adjustment from wet to dry conditions in the field, indicating phenotypic plasticity and importantly, the ability to increase hydraulic safety in the short term. In the nursery, seedlings from drier populations had smaller leaves and a lower leaf hydraulic vulnerability, suggesting that key traits associated with hydraulic safety are under strong genetic control. Overall, our study suggests a strong genetic control over traits associated with hydraulic safety, which may compromise the survival of wet-origin populations in drier future climates. However, phenotypic plasticity in physiological and morphological traits may confer sufficient hydraulic safety to facilitate genetic adaptation.