Loss Of Hydraulic Conductivity Research Articles

Stem notching decreases stem hydraulic conductance but does not influence drought impacts and post-drought recovery in Scots pine and Norway spruce.

The tight connection between the deterioration of xylem function and plant mortality under drought is well recognized. However, a lack of mechanistic understanding of how substantial conductivity loss influences plant performance under drought and during post-drought recovery hinders our ability to model tree responses to drought stress. We artificially induced a loss of 50% of xylem conducting area in Scots pine and Norway spruce saplings by stem notching and investigated plant performance under drought and during post-drought recovery. Plant mortality, xylem hydraulic conductivity, leaf water status and stomatal conductance were measured. We observed no preferential mortality of top plant parts (above the notches) compared to basal plant parts (below the notches), and no consistent trend in hydraulic conductivity loss was observed between top and basal parts of dying plants. Stem hydraulic conductivity, water status of the needles and stomatal conductance changed similarly between the top and basal parts during drought and post-drought recovery, which indicated the substantial hydraulic overcapacity of the stems. The recovery of stomatal conductance demonstrated prominent hysteresis due to non-hydraulic stomatal limitations. The results obtained are highly important for modelling the influence of plant hydraulic impairment on plant performance under drought and during post-drought recovery.

Stem hydraulic conductivity and embolism resistance of Quercus species are associated with their climatic niche.

The hydraulic traits of a plant species may reflect its climate adaptations. Southwest China is considered as a biodiversity hotpot of the genus Quercus (oak). However, the hydraulic adaptations of Asian oaks to their climate niches remain unclear. Ten common garden-grown oak species with distinct natural distributions in eastern Asia were used to determine their stem xylem embolism resistance (water potential at 50% loss of hydraulic conductivity, P50), stem hydraulic efficiency (vessel anatomy and sapwood specific hydraulic conductivity (Ks)) and leaf anatomical traits. We also compiled four key functional traits: wood density, hydraulic-weighted vessel diameter, Ks and P50 data for 31 oak species from previous literature. We analyzed the relationship between hydraulic traits and climatic factors over the native ranges of 41 oak species. Our results revealed that the 10 Asian oak species, which are mainly distributed in humid subtropical habitats, possessed a stem xylem with low embolism resistance and moderate hydraulic efficiency. The deciduous and evergreen species of the 10 Asian oaks differed in the stem and leaf traits related to hydraulic efficiency. Ks differed significantly between the two phenological groups (deciduous and evergreens) in the 41-oak dataset. No significant difference in P50 between the two groups was found for the 10 Asian oaks or the 41-oak dataset. The oak species that can distribute in arid habitats possessed a stem xylem with high embolism resistance. Ks negatively related to the humidity of the native range of the 10 Asian oaks, but showed no trend when assessing the entire global oak dataset. Our study suggests that stem hydraulic conductivity and embolism resistance in Quercus species are shaped by their climate niche. Our findings assist predictions of oak drought resistance with future climate changes for oak forest management.

The impact of climate change on growth and drought-induced mortality risk of Robinia pseudoacacia plantations along a precipitation gradient on the Chinese Loess Plateau

Robinia pseudoacacia has been widely planted on the Chinese Loess Plateau for soil and water conservation. Declining growth of R. pseudoacacia plantations has become a recently emerging challenge for revegetation and sustainable forest management in this region. In the suitable area of R. pseudoacacia, we selected four representative study sites along a precipitation gradient (aridity indexes of 1.3 at Fufeng, 1.5 at Changwu, 1.9 at Ansai, and 2.4 at Mizhi) and used an integrated Biome BioGeochemistry-plant hydraulic traits model (BBGC-SPERRY) to assess the climate change impacts on net primary productivity (NPP), maximum leaf area index (LAImax), actual evapotranspiration (AET), and annual average percentage loss of whole-plant hydraulic conductance (APLK) for R. pseudoacacia. We adopted climate scenario data derived from five downscaled Global Circulation Models under three Representative Concentration Pathways (RCPs2.6, RCP4.5 and RCP8.5) and set baseline (1981–2020), near future (2021–2060, NFP) and far future (2061–2099, FFP) periods. Our projections show annual precipitation will significantly increase during the NFP and FFP at the Ansai and Mizhi sites and increase slightly at the Fufeng and Changwu sites during the FFP; annual maximum and minimum temperatures will increase under all RCPs. The mean NPP and LAImax of R. pseudoacacia will significantly decrease by 9.9% (RCP4.5) to 49.1% (RCP2.6) during the NFP and by 7.4% (RCP4.5) to 50.1% (RCP2.6) during the FFP at the Changwu, Ansai, and Mizhi sites; however, both measures will significantly increase during the NFP and FFP at the Fufeng site. The AET will significantly increase during the NFP and FFP at all sites. The maximum APLK will increase from south to north along the precipitation gradient, with values greater than 60% at the Changwu, Ansai, and Mizhi sites during the NFP and FFP. These results indicate the drought-induced mortality risk of R. pseudoacacia will increase under future climate change.

Impact of freeze-thaw-induced pit aspiration on stem water transport in the subalpine conifer Abies veitchii.

During winter, subalpine conifers experience frequent freeze-thaw cycles in stem xylem that may cause embolism and pit aspiration due to increased water volume during the sap to ice transition. This study examined the occurrence and ecological impacts of a combination of freeze-thaw-induced pit aspiration and embolism triggered by natural and artificial stem freezing. In subalpine Veitch's fir (Abies veitchii) trees, the fraction of closed pits and embolized tracheids as well as conductivity losses were measured to examine pit aspiration and its effects. When trees incurred mild drought stress in February and early March, 70%-80% of stem conductivity was lost. Cryo-scanning electron microscopy indicated <20% embolized tracheids but ∼90% closed pits. Severe drought stress in late March caused 96% ± 1.2% (mean ± standard error) loss of stem conductivity, while the fraction of embolized tracheids increased to 64% ± 6.6%, and aspirated pit fraction decreased to 23% ± 5.6%. Experimental freeze-thaw cycles also increased pit aspiration from 7.1% ± 0.89% to 49% ± 10%, and the fraction of closed pits was positively correlated to the percent loss of stem hydraulic conductivity. The results indicated that freezing-induced pit aspiration is an important factor for stem xylem dysfunction under mild drought, and upon severe drought in winter; stem water transport is predominantly inhibited by xylem embolism.

Nitrogen deposition increases xylem hydraulic sensitivity but decreases stomatal sensitivity to water potential in two temperate deciduous tree species

Although the effects of nitrogen deposition on tree water relations are studied extensively, its impact on the relative sensitivities of stomatal and xylem hydraulic conductance to vapor pressure deficit and water potential is still poorly understood. This study investigated the effects of a 7-year N deposition treatment on the responses of leaf water relations and sensitivity of canopy stomatal conductance to vapor pressure deficit (VPD) and water potential, as well as the sensitivity of branch hydraulic conductance to water potential in a dominant tree species (Quercus wutaishanica) and an associated tree species (Acer mono) in a temperate forest. It was found that the N deposition increased stomatal sensitivity to VPD, decreased stomatal sensitivity to water potential, and increased the vulnerability of the hydraulic system to cavitation in both species. The standardized stomatal sensitivity to VPD, however, was not affected by the N deposition, indicating that the stomata maintained the ability to regulate the water balance under nitrogen deposition condition. Although the increased stomatal sensitivity to VPD could compensate the decreased stomatal sensitivity to water potential to some extent, the combined response would increase the percentage loss of hydraulic conductivity (PLC) when 50 % loss in stomatal conductance occurred, particularly in the dominant species Q. wutaishanica. The result indicates that N deposition would increase the risk of hydraulic failure in those species if the soil and/or air becomes drier under future climate change scenarios. The results of the study can have significant implications on the modelling of ecosystem vulnerability to drought under the scenario of atmospheric nitrogen deposition.

Coordination of leaf hydraulic and economic traits in Cinnamomum camphora under impervious pavement

BackgroundPaved urban environments can pose great threats to the physiological functioning and ecological services of street trees. In this context, assessment of leaf phenotypic plasticity is crucial for understanding the ecological strategy of tree species under impervious pavements.ResultsIn this study, we measured a set of leaf economic traits, hydraulic traits of Cinnamomum camphora, and surrounding environmental factors in a street site (the soil was covered by the impervious pavement) and a park site (the soil was covered by grass) in Hefei, eastern China. Compared with the park site, trees in the street site had higher stomatal length (SL), leaf thickness (LT), maximum photochemical quantum yield of photosystem II (Y(II)), and lower stomatal density (SD), specific leaf area (SLA), the leaf water potential at 50% loss of hydraulic conductance (P50), and leaf turgor loss point (TLP). Redundancy analysis showed that air relative humidity and volumetric soil water content caused these traits to be altered.ConclusionsOur results showed that C. camphora adapted to the street pavement environment through the coordination of leaf economic and leaf hydraulic traits, and adopted the slow investment return type in the leaf economic spectrum and high drought resistance to meet its actual physiological needs. This finding provides a new perspective for understanding the physiological strategies of street trees to adapt to urban pavement environments.

Predicting resilience through the lens of competing adjustments to vegetation function

There is a pressing need to better understand ecosystem resilience to droughts and heatwaves. Eco-evolutionary optimization approaches have been proposed as means to build this understanding in land surface models and improve their predictive capability, but competing approaches are yet to be tested together. Here, we coupled approaches that optimize canopy gas exchange and leaf nitrogen investment, respectively, extending both approaches to account for hydraulic impairment. We assessed model predictions using observations from a native Eucalyptus woodland that experienced repeated droughts and heatwaves between 2013 and 2020, whilst exposed to an elevated [CO2 ] treatment. Our combined approaches improved predictions of transpiration and enhanced the simulated magnitude of the CO2 fertilization effect on gross primary productivity. The competing approaches also worked consistently along axes of change in soil moisture, leaf area, and [CO2 ]. Despite predictions of a significant percentage loss of hydraulic conductivity due to embolism (PLC) in 2013, 2014, 2016, and 2017 (99th percentile PLC > 45%), simulated hydraulic legacy effects were small and short-lived (2 months). Our analysis suggests that leaf shedding and/or suppressed foliage growth formed a strategy to mitigate drought risk. Accounting for foliage responses to water availability has the potential to improve model predictions of ecosystem resilience.

The Role of Hydraulic Failure in a Massive Mangrove Die-Off Event.

Between late 2015 and early 2016, more than 7,000 ha of mangrove forest died along the coastline of the Gulf of Carpentaria, in northern Australia. This massive die-off was preceded by a strong 2015/2016 El Niño event, resulting in lower precipitation, a drop in sea level and higher than average temperatures in northern Australia. In this study, we investigated the role of hydraulic failure in the mortality and recovery of the dominant species, Avicennia marina, 2 years after the mortality event. We measured predawn water potential (Ψpd) and percent loss of stem hydraulic conductivity (PLC) in surviving individuals across a gradient of impact. We also assessed the vulnerability to drought-induced embolism (Ψ50) for the species. Areas with severe canopy dieback had higher native PLC (39%) than minimally impacted areas (6%), suggesting that hydraulic recovery was ongoing. The high resistance of A. marina to water-stress-induced embolism (Ψ50 = −9.6 MPa), indicates that severe water stress (Ψpd < −10 MPa) would have been required to cause mortality in this species. Our data indicate that the natural gradient of water-stress enhanced the impact of El Niño, leading to hydraulic failure and mortality in A. marina growing on severely impacted (SI) zones. It is likely that lowered sea levels and less frequent inundation by seawater, combined with lower inputs of fresh water, high evaporative demand and high temperatures, led to the development of hyper-salinity and extreme water stress during the 2015/16 summer.

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

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. 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. 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. 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.

Lianas and Trees From a Seasonally Dry and a Wet Tropical Forest Did Not Differ in Embolism Resistance but Did Differ in Xylem Anatomical Traits in the Dry Forest

One of the most prominent changes in neotropical forests has been the increase in abundance and size of lianas. Studies suggest that lianas have more acquisitive strategies than trees, which could allow them to take advantage of water more effectively when it is available in water-limited forests, but few studies compared across growth form (i.e., lianas vs. trees) and forest type (i.e., wet vs. seasonally dry). We measured hydraulic and anatomical traits of co-occurring lianas and trees that convey drought resistance (xylem embolism resistance and intervessel pit membranes) and water transport capacity (xylem vessel diameter and density) in a seasonally dry and a wet evergreen tropical forest to address: (1) Are there differences between vulnerability to embolisms (P50—water potential at 50% loss of hydraulic conductivity) and hydraulic safety margins (HSM) across growth form and forest type? (2) How do vessel diameter and density vary across growth form and forest type? (3) Are there differences in xylem intervessel pit membrane thickness across growth form and forest type and does it predict xylem embolism vulnerability in trees and lianas? We examined hydraulic and xylem anatomical traits of 32 species—eight lianas and eight trees in each forest type. We found no difference in P50and HSMs between lianas and trees and between the wetter and drier forest. Dry forest lianas had 81% greater maximum vessel diameter and 125% greater range in vessel diameter sizes than dry forest trees but, there was no significant difference between life forms in the wet forest. Dry forest species had 50% greater vessel density and 30% greater maximum pit membrane thickness than wet forest ones. Maximum pit membrane thickness was correlated to P50and HSMs. The main difference between lianas and trees occurred in the dry forest, where lianas had larger maximum xylem vessel size than trees, implying that they have proportionally greater hydraulic conductive capacity than the trees in seasonal forests.

Pathogen-induced hydraulic decline limits photosynthesis and starch storage in grapevines (Vitis sp.).

Xylella fastidiosa (Xf) is the bacterial pathogen responsible for Pierce's Disease (PD) in grapevine (Vitis vinifera L.) and numerous diseases in agriculturally and ecologically important species. Current theory suggests that localized inoculations via insect feeding lead to bacterial spread through the xylem, reducing water transport capacity, leading to declines in productivity, and ultimately death. Yet, the underlying mechanisms of Xf-induced mortality are not fully understood. In this study, we documented the development of PD symptoms over 12-13 weeks postinoculation. Subsequently assessed photosynthetic capacity, starch storage, and stem hydraulics in four grapevine genotypes (two PD-resistant and two PD-susceptible), comparing those physiological changes to control plants. PD-susceptible genotypes showed a coordinated decline in photosynthesis, starch storage, and stem hydraulics, whereas Xf-inoculation led to no change in starch and stem hydraulics in the PD-resistant genotypes. Together these data support the idea of a link between loss of hydraulic conductivity due to tylosis production with a downstream photosynthetic decline and starch depletion in the PD-susceptible genotypes. Our data support the theory that hydraulic failure and carbon starvation underlie plant mortality resulting from PD.

Lack of hydraulic recovery as a cause of post-drought foliage reduction and canopy decline in European beech.

Summary European beech (Fagus sylvatica) was among the most affected tree species during the severe 2018 European drought. It not only suffered from instant physiological stress but also showed severe symptoms of defoliation and canopy decline in the following year.To explore the underlying mechanisms, we used the Swiss‐Canopy‐Crane II site and studied in branches of healthy and symptomatic trees the repair of hydraulic function and concentration of carbohydrates during the 2018 drought and in 2019.We found loss of hydraulic conductance in 2018, which did not recover in 2019 in trees that developed defoliation symptoms in the year after drought. Reduced branch foliation in symptomatic trees was associated with a gradual decline in wood starch concentration throughout summer 2019. Visualization of water transport in healthy and symptomatic branches in the year after the drought confirmed the close relationship between xylem functionality and supported branch leaf area.Our findings showed that embolized xylem does not regain function in the season following a drought and that sustained branch hydraulic dysfunction is counterbalanced by the reduction in supported leaf area. It suggests acclimation of leaf development after drought to mitigate disturbances in canopy hydraulic function.

Detection of acoustic events in lavender for measuring xylem vulnerability to embolism and cellular damage.

Acoustic emission analysis is promising to investigate the physiological events leading to drought-induced injury and mortality. However, their nature and source are not fully understood, making this technique difficult to use as a direct measure of the loss of xylem hydraulic conductance. Acoustic emissions were recorded during severe dehydration in lavender plants (Lavandula angustifolia) and compared with the dynamics of embolism development and cell damage. The timing and characteristics of acoustic signals from two independent recording systems were compared by principal component analysis (PCA). Changes in water potential, branch diameter, loss of hydraulic conductance, and cellular damage were also measured to quantify drought-induced damages. Two distinct phases of acoustic emissions were observed during dehydration: the first one associated with a rapid loss of diameter and a significant increase in loss of xylem conductance (90%), and the second with slower changes in diameter and a significant increase in cellular damage. Based on PCA, a developed algorithm discriminated hydraulic-related acoustic signals from other sources, proposing a reconstruction of hydraulic vulnerability curves. Cellular damage preceded by hydraulic failure seems to lead to a lack of recovery. The second acoustic phase would allow detection of plant mortality.

No Evidence for Light-Induced Embolism Repair in Cut Stems of Drought-Resistant Mediterranean Species under Soaking.

(1) Recent studies suggested that stem photosynthesis could favor bark water uptake and embolism recovery when stem segments are soaked in water under light conditions, but evidence for this phenomenon in drought-resistant Mediterranean species with photosynthetic stems is missing. (2) Embolism recovery upon immersion in water for 2 h–4 h under light was assessed (i) via a classical hydraulic method in leafless Fraxinus ornus and Olea europaea branch segments stressed to xylem water potentials (Ψxyl) inducing ca. 50% loss of hydraulic conductivity (PLC) and (ii) via X-ray micro-CT imaging of the stem segments of drought-stressed potted F. ornus saplings. Hydraulic recovery was also assessed in vivo in intact drought-stressed F. ornus saplings upon soil re-irrigation. (3) Intact F. ornus plants recovered hydraulic function through root water uptake. Conversely, the soaked stem segments of both species did not refill embolized conduits, although Ψxyl recovered to pre-stress levels (between −0.5 MPa and −0.2 MPa). (4) We hypothesize that xylem embolism recovery through bark water uptake, even in light conditions, may not be a common phenomenon in woody plants and/or that wounds caused by cutting short stem segments might inhibit the refilling process upon soaking.

Lack of phenotypic plasticity in leaf hydraulics for 10 woody species common to urban forests of North China.

The survival and performance of urban forests are increasingly challenged by urban drought, consequently compromising the sustainability and functionality of urban vegetation. Plant-water relations largely determine species drought tolerance, yet little is known about the hydraulics of urban forest species. Here, we report the leaf hydraulic and carbon traits that govern plant growth and drought resistance, including vulnerability to embolism, hydraulic conductivity and leaf gas exchange characteristics, as well as morphological traits that are potentially linked with these physiological attributes, with the aim of guiding species selection and management in urban forests. Plant materials were collected from mature shrubs and trees on our university campus in Beijing, representing 10 woody species common to urban forests in north China. We found that the leaf embolism resistance, represented by the water potential inducing 50% loss of hydraulic conductivity (P50), as well as the hydraulic safety margin (HSM) defined by P50 and the water potential threshold at the inception of embolism (P12), varied remarkably across species, but was unrelated to growth form. Likewise, stem and leaf-specific hydraulic conductivity (Kstem and kl) was also highly species-specific. Leaf P50 was positively correlated with hydraulic conductivity. However, neither P50 nor hydraulic conductivity was correlated with leaf gas exchange traits, including maximum photosynthetic rate (Amax) and stomatal conductance (gs). Plant morphological and physiological traits were not related, except for specific leaf area, which showed a negative relationship with HSM. Traits influencing plant-water transport were primarily correlated with the mean annual precipitation of species climatic niche. Overall, current common woody species in urban forest environments differed widely in their drought resistance and did not have the capacity to modify these characteristics in response to a changing climate. Species morphology provides limited information regarding physiological drought resistance. Thus, screening urban forest species based on plant physiology is essential to sustain the ecological services of urban forests.

Testing the limits of plant drought stress and subsequent recovery in four provenances of a widely distributed subtropical tree species.

Drought-induced tree mortality may increase with ongoing climate change. Unraveling the links between stem hydraulics and mortality thresholds, and the effects of intraspecific variation, remain important unresolved issues. We conducted a water manipulation experiment in a rain-out shelter, using four provenances of Schima superba originating from a gradient of annual precipitation (1124-1796 mm) and temperature (16.4-22.4°C). Seedlings were droughted to three levels of percentage loss of hydraulic conductivity (i.e., P50 , P88 and P99) and subsequently rewatered to field capacity for 30 days; traits related to water and carbon relations were measured. The lethal water potential associated with incipient mortality was between P50 and P88 . Seedlings exhibited similar drought responses in xylem water potential, hydraulic conductivityand gas exchange. Upon rehydration, patterns of gas exchange differed among provenances but were not related to the climate at the origin. The four provenances exhibited a similar degree of stem hydraulic recovery, which was correlated with the magnitude of antecedent drought and stem soluble sugar at the end of the drought. Results suggest that there were intraspecific differences in the capacity of S. superba seedlings for carbon assimilation during recovery, indicating a decoupling between gas exchange recovery and stem hydraulics across provenances.

Stem and leaf xylem of angiosperm trees experiences minimal embolism in temperate forests during two consecutive summers with moderate drought.

Drought events may increase the likelihood that the plant water transport system becomes interrupted by embolism. Yet our knowledge about the temporal frequency of xylem embolism in the field is frequently lacking, as it requires detailed, long-term measurements. We measured xylem embolism resistance and midday xylem water potentials during the consecutive summers of 2019 and 2020 to estimate maximum levels of embolism in leaf and stem xylem of ten temperate angiosperm tree species. We also studied vessel and pit membrane characteristics based on light and electron microscopy to corroborate potential differences in embolism resistance between leaves and stems. Apart from A. pseudoplatanus and Q. petraea, eight species experienced minimum xylem water potentials that were close to or below those required to initiate embolism. Water potentials corresponding to ca. 12% loss of hydraulic conductivity (PLC) could occur in six species, while considerable levels of embolism around 50% PLC were limited to B. pendula and C. avellana. There was a general agreement in embolism resistance between stems and leaves, with leaves being equally or more resistant than stems. Also, xylem embolism resistance was significantly correlated to intervessel pit membrane thickness (TPM ) for stems, but not to vessel diameter and total intervessel pit membrane surface area of a vessel. Our data indicate that low amounts of embolism occur in most species during moderate summer drought, and that considerable levels of embolism are uncommon. Moreover, our experimental and TPM data show that leaf xylem is generally no more vulnerable than stem xylem.

Physiological and Biochemical Dynamics of Pinus massoniana Lamb. Seedlings under Extreme Drought Stress and during Recovery

In recent years, global forests have been facing an increase in tree mortality owing to increasing droughts. However, the capacity for plants to adjust their physiology and biochemistry during extreme drought and subsequent recovery is still unclear. Here, we used 1.5-year-old Pinus massoniana Lamb. seedlings and simulated drought conditions to achieve three target stress levels (50%, 85%, and 100% loss of stem hydraulic conductivity (PLC)), followed by rehydration. Needle water status, gas exchange, and biochemical parameters were assessed during drought and recovery. The results showed that drought had significantly negative impacts on needle water status and gas exchange parameters, with gas exchange declining to 0 after PLC85 was achieved. Soluble protein concentration (SPC), soluble sugar concentration (SSC), malondialdehyde (MDA) content, superoxide dismutase (SOD) activity, and needle water-use efficiency showed fluctuations. The activity of antioxidant enzymes and the values of osmotic regulators were then gradually decreased as the physiological and biochemical functions of seedlings were disturbed. Seedlings showed a stronger ability to recover from PLC50 than PLC85 and PLC100. We conclude that the physiological and biochemical recovery of P. massoniana seedlings is more likely to be inhibited when plants experience increasing drought stress that induces 85% and greater loss of hydraulic conductance.

Contrast in vulnerability to freezing-induced xylem embolism contributes to divergence in spring phenology between diffuse- and ring-porous temperate trees

The spring phenology and growth strategy of temperate tree species can be strongly linked to their sensitivity to frosts, which deserve more profound investigations under the background of climate warming particularly considering the advancement of spring phenology as well as the increase in frequency and intensity of spring cold waves. Spring phenologies, stem radial growth characteristics, frost sensitivity of leaves and stem hydraulic systems were studied in five diffuse-porous and five ring-porous temperate tree species under a common garden condition. The results showed that the spring leaf phenology of the diffuse-porous species was one to two weeks earlier than that of the ring-porous species. The ring-porous species had significantly higher stem hydraulic conductivity than the diffuse-porous species (1.81 and 0.95 kg·m −1 ·s −1 ·MPa −1 , P < 0.05) but were more vulnerable to freeze-thaw induced xylem embolism than the latter. After a simulated freeze-thaw event, the average percentage loss of hydraulic conductivity in the current year shoots increased from 26.0% (native embolism) to 86.7% in the ring-porous species, while it only increased from 21.3% to 38.3% in the diffuse-porous species. The spring phenology was clearly correlated with vulnerability to freeze-thaw induced embolism, with the more vulnerable ring-porous species exhibited substantially delayed phenology to reduce risks of catastrophic hydraulic dysfunction during spring frosts. Nevertheless, ring-porous species can offset the postponed onset of growth and gained even higher annual growth due to significantly higher hydraulic efficiency and leaf gas exchange rates. Contrasts between ring-porous and diffuse-porous species in resistance to freeze-thaw induced embolism suggest that they face different selective pressures from early spring frosts, which may at least be partially responsible for their divergence in spring phenology and growth strategy and can potentially lead to different responses to climate regime shifts.

Hydraulic traits are coupled with plant anatomical traits under drought–rewatering cycles in Ginkgo biloba L.

Investigating the responses of plant anatomical traits of trees to drought-rewatering cycles helps us to understand their responses to climate change; however, such work has not been adequately reported. In this study, Ginkgo biloba L. saplings were subjected to moderate, severe, extreme and lethal drought conditions by withholding water according to the percentage loss of hydraulic conductivity (PLC) and rewatering on a regular basis. Samples of phloem, cambium and xylem were collected to quantify their cellular properties including cambium and phloem cell vitality, xylem growth ring width, pit aspiration rates and pit membrane thickness using light microscopy and transmission microscopy. The results showed that the mortality rate of G. biloba saplings reached 90% at approximately P88 (xylem water potential inducing 88% loss of hydraulic conductivity). The onset of cambium and phloem cell mortality might be in accordance with that of xylem embolism. Close negative correlations between xylem water potential and PLC and between xylem water potential and cambium and phloem mortality suggested that xylem hydraulic traits are coupled with anatomical traits under declining xylem water potential. Cambium and phloem cell vitality as well as xylem growth ring width decreased significantly with increasing drought conditions. However, xylem pit membrane thickness, cambial zone width and cambial cell geometry were not affected by the drought-rewatering cycles. The tracheid radial diameter, intertracheid cell wall thickness and tracheid density decreased significantly during both drought conditions and rewatering conditions. In addition to hydraulic traits, cambium and phloem cell vitality can be used as anatomical traits to evaluate the mortality of G. biloba under drought. Future work is proposed to observe the dynamics of pit aspiration rates under drought-rewatering cycles in situ to deepen our understanding of the essential role of bordered pits in the 'air-seeding' mechanism.