Hydraulic properties and drought response of a tropical bamboo (Cephalostachyum pergracile).

Bamboo plants are an essential component of tropical ecosystems, yet their vulnerability to climate extremes, such as drought, is poorly understood due to limited knowledge of their hydraulic properties. Cephalostachyum pergracile, a commonly used tropical bamboo species, exhibited a substantially higher mortality rate than other co-occurring bamboos during a severe drought event in 2019, but the underlying mechanisms remain unclear. This study investigated the leaf and stem hydraulic traits related to drought responses, including leaf-stem embolism resistance (P50leaf; P50stem) estimated using optical and X-ray microtomography methods, leaf pressure-volume and water-releasing curves. Additionally, we investigated the seasonal water potentials, native embolism level (PLC) and xylem water source using stable isotope. We found that C. pergracile exhibited strong resistance to embolism, showing low P50leaf, P50stem, and turgor loss point, despite its rapid leaf water loss. Interestingly, its leaves displayed greater resistance to embolism than its stem, suggesting a lack of effective hydraulic vulnerability segmentation (HVS) to protect the stem from excessive xylem tension. During the dry season, approximately 49% of the water was absorbed from the upper 20-cm-deep soil layer. Consequently, significant diurnal variation in leaf water potentials and an increase in midday PLC from 5.87 ± 2.33% in the wet season to 12.87 ± 4.09% in the dry season were observed. In summary, this study demonstrated that the rapid leaf water loss, high reliance on surface water, and a lack of effective HVS in C. pergracile accelerated water depletion and increased xylem embolism even in the typical dry season, which may explain its high mortality rate during extreme drought events in 2019.

Sources and consequences of mismatch between leaf disc and whole-leaf leaf mass per area (LMA).

PREMISE: Leaf mass per area (LMA), which is an important functional trait in leaf economic spectrum and plant growth analysis, is measured from leaf discs or whole leaves. Differences between the measurement methods may lead to large differences in the estimates of LMA values. METHODS: We examined to what extent estimates of LMA based on whole leaves match those based on discs using 334 woody species from a wide range of biomes (tropics, subtropics, savanna, and temperate), whether the relationship varied by leaf morphology (tissue density, leaf area, leaf thickness), punch size (0.6- and 1.0-cm diameter), and whether the extent of intraspecifc variation for each species matches. RESULTS: Disc-based estimates of species mean LMA matched the whole-leaf estimates well, and whole-leaf LMA tended to be 9.69% higher than leaf-disc LMA. The ratio of whole-leaf LMA to leaf-disc LMA was higher for species with higher leaf tissue density and larger leaves, and variance in the ratio was greater for species with lower leaf tissue density and thinner leaves. Estimates based on small leaf discs also inflated the ratio. The extent of the intraspecific variation only weakly matched between whole-leaf and disc-based estimates (R2 = 0.08). CONCLUSIONS: Our results suggest that simple conversion between whole-leaf and leaf-disc LMA is difficult for species obtained with a small leaf punch, but it should be possible for species obtained with a large+ leaf punch. Accurately representing leaf traits will likely require careful selection between leaf-disc and whole-leaf traits depending on the objectives. Quantifying intraspecific variation using leaf discs should be also considered with caution.

Response of four evergreen savanna shrubs to an incidence of extreme drought: high embolism resistance, branch shedding and maintenance of nonstructural carbohydrates.

Extreme drought events are becoming frequent globally, resulting in widespread plant mortality and forest dieback. Although savanna vegetation cover ~20% of the earth's land area, their responses to extreme drought have been less studied than that of forests. Herein, we quantified branch dieback, individual mortality and the associated physiological responses of four evergreen shrubs (Tarenna depauperate Hutch., Maytenus esquirolii (H. Lév.) C.Y. Cheng, Murraya exotica L., Jasminum nudiflorum Lindl.) in a savanna ecosystem in Southwest China to an incidence of extreme drought during 2019 and 2020. We found that 80-100% of the individuals of these species exhibited branch dieback, whereas individual mortality was only found in T. depauperate (4.5%). All species showed high resistance to stem embolism (P50, water potential at 50% loss of hydraulic conductivity ranged from -5.62 to -8.6 MPa), whereas the stem minimum water potentials reached -7.6 to ca -10.0 MPa during the drought. The low water potential caused high native embolism levels (percentage loss of hydraulic conductivity (PLC) 23-65%) in terminal branches, and the remaining stems maintained 15-35% PLC at the end of the drought. Large within-individual variations in stem vulnerability to embolism were detected, and shedding of vulnerable branches could be a mechanism for shrubs to reduce water and carbon consumption. Overall, the content of total nonstructural carbohydrates (NSC) and their components in the stem were generally comparable to or higher than those in the rainy season in three of the four species. Because the leaves were turgor-less for most time during the drought, high NSC levels during the drought could be due to recycling of NSC from dead branches or translocation from roots. Our results suggest high tolerance of savanna shrub species to extreme drought, which could be facilitated by high embolism resistance in some stems and shedding of vulnerable branches to maintain individual water and carbon balance.

Hydraulic prediction of drought-induced plant dieback and top-kill depends on leaf habit and growth form.

Hydraulic failure caused by severe drought contributes to aboveground dieback and whole-plant death. The extent to which dieback or whole-plant death can be predicted by plant hydraulic traits has rarely been tested among species with different leaf habits and/or growth forms. We investigated 19 hydraulic traits in 40 woody species in a tropical savanna and their potential correlations with drought response during an extreme drought event during the El Niño-Southern Oscillation in 2015. Plant hydraulic trait variation was partitioned substantially by leaf habit but not growth form along a trade-off axis between traits that support drought tolerance versus avoidance. Semi-deciduous species and shrubs had the highest branch dieback and top-kill (complete aboveground death) among the leaf habits or growth forms. Dieback and top-kill were well explained by combining hydraulic traits with leaf habit and growth form, suggesting integrating life history traits with hydraulic traits will yield better predictions.

Quantifying vulnerability to embolism in tropical trees and lianas using five methods: can discrepancies be explained by xylem structural traits?

Vulnerability curves (VCs) describe the loss of hydraulic conductance against increasing xylem tension, providing valuable insights about the response of plant water transport to water stress. Techniques to construct VCs have been developed and modified continuously, but controversies continue. We compared VCs constructed using the bench-top dehydration (BD), air-injection-flow (AI), pneumatic-air-discharge (PAD), optical (OP) and X-ray-computed microtomography (MicroCT) methods for tropical trees and lianas with contrasting vessel lengths. The PAD method generated highly vulnerable VCs, the AI method intermediate VCs, whereas the BD, OP and MicroCT methods produced comparable and more resistant VCs. Vessel-length and diameter accounted for the overestimation ratio of vulnerability estimated using the AI but not the PAD method. Compared with directly measured midday embolism levels, the PAD and AI methods substantially overestimated embolism, whereas the BD, MicroCT and OP methods provided more reasonable estimations. Cut-open vessels, uncertainties in maximum air volume estimations, sample-length effects, tissue cracks and shrinkage together may impede the reliability of the PAD method. In conclusion, we validate the BD, OP and MicroCT methods for tropical plants, whereas the PAD and AI need further mechanistic testing. Therefore, applications of VCs in estimating plant responses to drought need to be cautious.

Vessel-length determination using silicone and air injection: are there artifacts?

Xylem vessels are used by most angiosperm plants for long-distance water and nutrient transport. Vessel length is one of the key functional traits determining plant water-transport efficiency. Additionally, determination of maximum vessel length is necessary for correct sample collection and measurements in hydraulic studies to avoid open-vessel and cutting-under-tension artifacts. Air injection and silicone injection (BLUESIL RTV141A and B mixtures) are two widely used methods for maximum vessel length determination. However, the validity of both methods needs to be carefully tested for species with different vessel lengths. In this study, we tested the air-injection and silicone-injection methods using eight species with different vessel lengths: short (<0.5 m), medium (0.5-1 m) and long (>1 m). We employed a novel approach using RTV141A injection without the RTV141B hardener as a reference method because RTV141A cannot penetrate inter-vessel pit membranes and is not prone to hardening/solidification effects during the injection process. The results revealed that the silicone-injection method substantially underestimated the maximum vessel length of all eight species. However, the air-injection method tended to overestimate the maximum vessel length in five out of eight species. The ratio of underestimation of the silicone-injection method was higher for species with longer vessels, but the overestimation of the air-injection method was independent of the vessel length. Moreover, air injection with different pressures-ranging from 40 to 300 kPa-resulted in comparable results. We conclude that the conventional silicone-injection method can underestimate the vessel length, whereas the air-injection method can overestimate the maximum vessel length, particularly for long-vessel led species. We recommend RTV141A-only injection for determining the maximum vessel length, and it can also be used to validate the use of the air-injection and conventional silicone-injection methods for a given species.