A unit pipe pneumatic model to simulate gas kinetics during measurements of embolism in excised angiosperm xylem.

The pneumatic method has been introduced to quantify embolism resistance in plant xylem of various organs by applying a partial vacuum to cut-open xylem. Despite the similarity in vulnerability curves between the pneumatic and other methods, a modeling approach is needed to investigate if changes in xylem embolism during dehydration can be accurately quantified based on gas diffusion kinetics. Therefore, a unit pipe pneumatic (UPPn) model was developed to estimate gas extraction from intact conduits, which were axially interconnected by inter-conduit pit membranes to cut-open conduits. The physical laws used included Fick's law for diffusion, Henry's law for gas concentration partitioning between liquid and gas phases at equilibrium and the ideal gas law. The UPPn model showed that 91% of the extracted gas came from the first five series of embolized, intact conduits and only 9% from the aqueous phase after 15 s of simulation. Considering alternative gas sources, embolism resistance measured with a pneumatron device was systematically overestimated by 2-17%, which corresponded to a typical measuring error of 0.11 MPa for P50 (the water potential equivalent to 50% of the maximum amount of gas extracted). It is concluded that pneumatic vulnerability curves directly measure embolism of intact conduits due to the fast movement of gas across interconduit pit membranes, while gas extraction from sap and diffusion across hydrated cell walls is about 100 times slower. We expect that the UPPn model will also contribute to the understanding of embolism propagation based on temporal gas dynamics.

The theory behind vessel length determination using gas flow rates and comparison between two pneumatic methods based on seven woody species.

In plants, xylem vessel length is important for long-distance water transport; however, the currently used methods for vessel length measurement are inconvenient and time-consuming. The recently developed semi-automated Pneumatron is a device based on the pneumatic theory that is similar to the air-injection method, and can rapidly estimate vessel length. Mean vessel length was compared between the Pneumatron and the air-injection method in seven woody species with a wide range of vessel lengths (2.3-78.7 cm). The results were consistent between the two methods, regardless of whether the same or different samples were used. The theory underlying the gas flow in vessels was improved and expanded, and compared to that underlying the water flow in order to better understand the pneumatic processes within a stem sample. Moreover, a new and simple equation for gas flow in vessels was derived based on the molar gas flow (mol s-1) rather than volume flow, because the former remains constant with distance throughout the stem axis. We strongly recommend using the Pneumatron in future studies owing to its low cost, convenience, rapidity, and simple operation. However, a number of potential issues need to be considered to avoid artifacts during measurements.

Leaf Venation Architecture in Relation to Leaf Size Across Leaf Habits and Vein Types in Subtropical Woody Plants.

Leaves are enormously diverse in their size and venation architecture, both of which are core determinants of plant adaptation to environments. Leaf size is an important determinant of leaf function and ecological strategy, while leaf venation, the main structure for support and transport, determines the growth, development, and performance of a leaf. The scaling relationship between venation architecture and leaf size has been explored, but the relationship within a community and its potential variations among species with different vein types and leaf habits have not been investigated. Here, we measured vein traits and leaf size across 39 broad-leaved woody species within a subtropical forest community in China and analyzed the scaling relationship using ordinary least squares and standard major axis method. Then, we compared our results with the global dataset. The major vein density, and the ratio of major (1° and 2°) to minor (3° and higher) vein density both geometrically declined with leaf size across different vein types and leaf habits. Further, palmate-veined species have higher major vein density and a higher ratio of major to minor vein density at the given leaf size than pinnate-veined species, while evergreen and deciduous species showed no difference. These robust trends were confirmed by reanalyzing the global dataset using the same major vein classification as ours. We also found a tradeoff between the cell wall mass per vein length of the major vein and the major vein density. These vein scaling relationships have important implications on the optimization of leaf size, niche differentiation of coexisting species, plant drought tolerance, and species distribution.

The hydraulic architecture of an arborescent monocot: ontogeny-related adjustments in vessel size and leaf area compensate for increased resistance.

Bamboos are arborescent monocotyledons that have no secondary growth, but can continually produce conduits with diameters appropriate to the current size of the plant. Here, we studied bamboo hydraulic architecture to address the mechanisms involved in compensating for the increase in hydraulic resistance during ontogeny. We measured the hydraulic weighted vessel diameters (Dh ) at different distances from the apex along the stem of Bambusa textilis. The hydraulic resistance of different components and individuals of different heights were quantified using the high-pressure flowmeter method. The Dh showed tip-to-base widening with a scaling exponent in the range of those reported for trees. Although theoretical hydraulic conductivity decreased from base-to-tip, leaf-specific conductivity did not change. Leaves contributed the most to the whole-shoot hydraulic resistance, followed by the leaf-bearing branches. Roots contributed c. 13% to whole-plant resistance. Interestingly, taller individuals showed lower whole-shoot resistance owing to an increased number of resistances in parallel (side-branches), while leaf-specific resistance was independent of plant size. Tip-to-base vessel widening and height-independent constant leaf-specific conductance could be mechanisms for hydraulic optimization in B. textilis. Similar patterns have also been found in woody plants with secondary growth, but this bamboo exhibits them without secondary growth.

Inferring the role of pit membranes in solute transport from solute exclusion studies in living conifer stems.

The functional role of pits between living and dead cells has been inferred from anatomical studies but amassing physiological evidence has been challenging. Centrifugation methods were used to strip water from xylem conduits, permitting a more quantitative gravimetric determination of the water and solid contents of cell walls than is possible by more traditional methods. Quantitative anatomical evidence was used to evaluate the water volume in xylem conduits and the water content of living cells. Quantitative perfusion of stems with polyethylene glycol of different molecular weight was used to determine the solute-free space. We measured the portioning of water and solute-free space among anatomical components in stems and demonstrated that lignin impeded the free movement of solutes with molecular weight >300. Hence, movement of large solutes from living cells to xylem conduits is necessarily confined to pit structures that permit transmembrane solute transport via primary walls without lignin. The functional role of pits was additionally indicated by combining data in this paper with previous studies of unusual osmotic relationships in woody species that lack pits between dead wood fibers and vessels. The absence of pits, combined with the evidence of exclusion of solutes of molecular weight >300, explains the unexpected osmotic properties.

An improved centrifuge method for determining water extraction curves and vulnerability curves in the long-vessel species Robinia pseudoacacia.

Significant improvements to the centrifuge water-extraction method of measuring the percentage loss volume of water (PLV) and corresponding vulnerability curves (VCs) are reported. Cochard and Sperry rotors are both incapable of measuring the VCs of species with long vessels because of premature embolism induced by hypothetical nanoparticles that can be drawn into segments during flow measurement. In contrast, water extraction pushes nanoparticles out of the sample. This study focuses on a long-vessel species, Robinia pseudoacacia, for which many VCs have been constructed by different methods, and the daily water relations have been quantified. PLV extraction curves have dual Weibull curves, and this paper focuses on the second Weibull curve because it involves the extraction of water from vessels, as proven by staining methods. We demonstrate an improved water extraction method after evaporation correction that has accuracy to within 0.5%, shows good agreement with two traditional methods that are slower and less accurate, and is immune to nanoparticle artefacts. Using Poiseuille's Law and the geometry of vessels, we argue that the percentage loss of conductivity (PLC) equals 2PLV-PLV2 in a special case where all vessels, regardless of size, have the same vulnerability curve. In this special case, this equation predicts the data reasonably well.

Drought inhibits photosynthetic capacity more in females than in males of Populus cathayana.

We investigated sex-related photosynthetic responses to drought in the dioecious species, Populus cathayana Rehd. Plants were subjected to two watering regimes (100% and 30% of field capacity) in a semi-controlled environment. Drought significantly decreased leaf area (LA), total number of leaves (TNL), specific leaf area (SLA), relative water content, net photosynthetic rate (P(n)), transpiration (E), stomatal conductance (g(s)), intercellular CO(2) concentration (C(i)), light saturation point (L(SP)), apparent quantum yield (Phi), carboxylation efficiency (CE), light-saturated photosynthetic rate (P(max)), maximum efficiency of PSII (F(v)/F(m)) and maximum effective quantum yield of PSII (Yield), and increased the total chlorophyll concentration (TC), CO(2) compensation point (Gamma), non-photochemical quenching coefficient, peroxidase (POD) activity and carbon isotope composition (delta(13)C). Moreover, differences between males and females were detected in many of these responses. In the drought treatment, males exhibited significantly higher LA, TNL, TC, concentration of carotenoids (Caro), P(n), E, g(s), C(i), L(SP), Phi, CE, P(max), F(v)/F(m), photochemical quenching coefficient, POD activity and delta(13)C, but a lower SLA, chlorophyll a/b ratio, carotenoids/total chlorophyll ratio and Gamma than females. However, Caro, L(SP), Gamma, Phi, CE and POD activity were apparently associated with sex-related resource demands, because significant differences in these traits were detected between the sexes under well-watered conditions. Our results indicate that drought stress limits photosynthetic capacity more in females than in males.