Haruhiko Taneda

The criteria for biomass partitioning of the current shoot: water transport versus mechanical support

In this study, we determine the theoretical criteria for biomass partitioning into the leaf and stem of the current shoot, using two quantitative models. The water transport model, based on the biochemical model of CO(2) assimilation, predicts the relationship between the water transport capacity per biomass investment in the stem (stem mass specific conductivity) and the partitioning of biomass that maximizes shoot productivity. The mechanical support model, based on Eulers buckling formula, predicts the relationship between the mechanical strength per biomass investment in the stem (the inverse relationship of stem mass density) and the partitioning of biomass to avoid mechanical failures such as lodging. These models predict the stem properties of mass specific conductivity and stem mass density that result in optimum partitioning just sufficient to provide adequate water transport and static mechanical support. In reality, the stem properties of plants differ from those predicted for optimum partitioning: the partitioning of biomass in the current shoot of both angiosperms and gymnosperms is mainly governed by the mechanical support criterion, although gymnosperms are probably more affected by the water transport criterion. This tendency is supported by actual measurements of biomass partitioning in plants.

Optimum leaf size predicted by a novel leaf energy balance model incorporating dependencies of photosynthesis on light and temperature

To clarify relationships between leaf size and the environment variables, we constructed an energy balance model for a single leaf incorporating Leuning’s stomatal conductance model and Farquhar’s leaf photosynthesis model. We ran this model for various environmental conditions paying particular attention to the leaf boundary layer. The leaf size maximizing the rate of photosynthesis per unit leaf area (A) at a high irradiance differed depending on the air temperature. In warm environments, A increased with decrease in leaf size, whereas in cool environments, there was the leaf size maximizing A. With the increase in leaf size, the CO2 concentration inside the leaf (Ci) decreased and the leaf temperature increased, both due to lower boundary layer conductance. At low air temperatures, the negative effect of low Ci on A in large leaves was compensated by the increase in leaf temperature towards the optimum temperature for A. This balance determined the optimum leaf size for A at low air temperatures. With respect to water use efficiency, large leaves tended to be advantageous, especially in cool environments at low-to-medium irradiances. Some temperature-dependent trends in leaf size observed in nature are discussed based on the present results.

Co-ordinated development of the leaf midrib xylem with the lamina in Nicotiana tabacum.

BACKGROUND AND AIMS The water-transport capacity of leaf venation is positively related to the leaf-lamina area, because the number and diameter of vein-xylem conduits are controlled to match the lamina area. This study aimed to investigate how this co-ordinated relationship between the leaf-lamina area and vein-xylem characteristics is achieved by examining the midrib xylem of tobacco leaves. METHODS The changes in the midrib-xylem characteristics over time were quantified using leaves with four different final lamina areas. The measured data were fitted to sigmoidal functions. From the constants of the fitted curves, the final values in mature leaves, maximal developmental rates (V(Dev)) and developmental duration (T(Dev)) were estimated for each of the xylem characteristics. Whether it is the lamina or the midrib xylem that drives the co-ordinated development was examined by lamina removal from unfolding leaves. The effects of the application of 0·1 % IAA (indole-3-acetic acid) to leaves with the laminas removed were also analysed. KEY RESULTS For both the leaf lamina and the midrib-xylem characteristics, the differences in final values among leaves with different lamina areas were more strongly associated with those in V(Dev). Notably, the V(Dev) values of the midrib-xylem characteristics were related to those of the leaf-lamina area. By lamina removal, the conduit diameter was reduced but the number of conduits did not significantly change. By IAA application, the decrease in the conduit diameter was halted, and the number of conduits in the midrib xylem increased. CONCLUSIONS According to the results, the V(Dev) values of the lamina area and the midrib-xylem characteristics changed in a co-ordinated manner, so that the water-transport capacity of the midrib xylem was positively related to the leaf-lamina area. The results also suggest that IAA derived from the leaf lamina plays a crucial role in the development of the leaf venation.

Leaf-lamina conductance contributes to an equal distribution of water delivery in current-year shoots of kudzu-vine shoot, Pueraria lobata

Leaf-lamina resistance, R(L), accounts for a large fraction of branch resistance across a wide range of plant species. This work hypothesized that large R(L) is essential for distributing water equally to leaves on the shoot, and tested this hypothesis through theoretical analyses and measurements using over 10-m-long current-year shoots of kudzu vine, Pueraria lobata [Willd.] Ohwi. First, the hydraulic architecture and the distribution of the motive force achieving equal distribution of water delivery were theoretically obtained by simulating water flow through a hypothetical shoot comprising an axial pathway and several lateral pathways as a stem and leaves, respectively, in a kudzu-vine shoot. The model predicts that large resistance of the lateral pathway relative to that of the axial pathway is associated strongly with small variation in the hydraulic conductance of a pathway from the base of the axial pathways to the lateral pathway among the nodes, rendering water delivery to each lateral pathway equal under small variation in motive force for water flow. For the kudzu-vine shoot, the measured ratio of the lateral (a petiole) to the axial (a stem) resistance was 115. When R(L) was added to the lateral pathway, the ratio increased to 1136. According to the model prediction, these values imply that the hydraulic conductance of a pathway comprising a stem and a petiole, K(BP), is favored strongly at the basal nodes, while the hydraulic conductance of a pathway including a stem, a petiole and a lamina, K(SL), is slightly different across the nodes. For the shoots with leaf lamina, the diurnal change in transpiration rate was not different between the leaves on the three nodes dividing the shoot into four parts. K(SL) was not related significantly to node number. Conversely, K(BP) at the distal node was ~0.06-fold that at the basal node. Furthermore, the motive force for water flow should vary by 6.64-fold among nodes to compensate for the favored distribution of K(BP), which is an unrealistic value. These results indicate that R(L) contributes largely to an equal distribution of water delivery in a shoot, supporting our hypothesis.

A theoretical approach to the relationship between wettability and surface microstructures of epidermal cells and structured cuticles of flower petals

BACKGROUND AND AIMS The epidermal surface of a flower petal is composed of convex cells covered with a structured cuticle, and the roughness of the surface is related to the wettability of the petal. If the surface remains wet for an excessive amount of time the attractiveness of the petal to floral visitors may be impaired, and adhesion of pathogens may be promoted. However, it remains unclear how the epidermal cells and structured cuticle contribute to surface wettability of a petal. METHODS By considering the additive effects of the epidermal cells and structured cuticle on petal wettability, a thermodynamic model was developed to predict the wetting mode and contact angle of a water droplet at a minimum free energy. Quantitative relationships between petal wettability and the geometries of the epidermal cells and the structured cuticle were then estimated. Measurements of contact angles and anatomical traits of petals were made on seven herbaceous species commonly found in alpine habitats in eastern Nepal, and the measured wettability values were compared with those predicted by the model using the measured geometries of the epidermal cells and structured cuticles. KEY RESULTS The model indicated that surface wettability depends on the height and interval between cuticular steps, and on a height-to-width ratio for epidermal cells if a thick hydrophobic cuticle layer covers the surface. For a petal epidermis consisting of lenticular cells, a repellent surface results when the cuticular step height is greater than 0·85 µm and the height-to-width ratio of the epidermal cells is greater than 0·3. For an epidermis consisting of papillate cells, a height-to-width ratio of greater than 1·1 produces a repellent surface. In contrast, if the surface is covered with a thin cuticle layer, the petal is highly wettable (hydrophilic) irrespective of the roughness of the surface. These predictions were supported by the measurements of petal wettability made on flowers of alpine species. CONCLUSIONS The results indicate that surface roughness caused by epidermal cells and a structured cuticle produces a wide range of petal wettability, and that this can be successfully modelled using a thermodynamic approach.

Structures of Bordered Pits Potentially Contributing to Isolation of a Refilled Vessel from Negative Xylem Pressure in Stems of Morus australis Poir.: Testing of the Pit Membrane Osmosis and Pit Valve Hypotheses.

Two hypotheses have been proposed to explain the mechanism preventing the refilling vessel water from being drained to the neighboring functional vessels under negative pressure. The pit membrane osmosis hypothesis proposes that the xylem parenchyma cells release polysaccharides that are impermeable to the intervessel pit membranes into the refilling vessel; this osmotically counteracts the negative pressure, thereby allowing the vessel to refill. The pit valve hypothesis proposes that gas trapped within intervessel bordered pits isolates the refilling vessel water from the surrounding functional vessels. Here, using the single-vessel method, we assessed these hypotheses in shoots of mulberry (Morus australis Poir.). First, we confirmed the occurrence of xylem refilling under negative pressure in the potted mulberry saplings. To examine the pit membrane osmosis hypothesis, we estimated the semi-permeability of pit membranes for molecules of various sizes and found that the pit membranes were not semi-permeable to polyethylene glycol of molecular mass <20,000. For the pit valve hypothesis, we formed pit valves in the intervessel pits in the short stem segments and measured the maximum liquid pressure up to which gases in bordered pits were retained. The threshold pressure ranged from 0.025 to 0.10 MPa. These values matched the theoretical value calculated from the geometry of the pit chamber (0.0692-0.101 MPa). Our results suggest that gas in the pits is retained by surface tension, even under substantial positive pressure to resolve gases in the refilling vessel, whereas the molecule size required for the pit membrane osmosis mechanism in mulberry would be unrealistically large.

Intra-specific trends of lumen and wall resistivities of vessels within the stem xylem vary among three woody plants

Water flow through xylem vessels encounters hydraulic resistance when passing through the vessel lumen and end wall. Comparative studies have reported that lumen and end wall resistivities co-limit water flow through stem xylem in several angiosperm woody species that have vessels of different average diameter and length. This study examined the intra-specific relationship between the lumen and end wall resistivities (Rlumen and Rwall) for vessels within the stem xylem using three deciduous angiosperm woody species found in temperate forest. Morus australis Poir. and Acer rufinerve Siebold et Zucc. are early- and late-successional species, and Vitis coignetiae Pulliat ex Planch is a woody liana. According to the Hagen-Poiseuille equation, Rlumen is proportional to the fourth power of vessel diameter (D), whereas vessel length (L) and inter-vessel pit area (Apit) determine Rwall. To estimate Rlumen and Rwall, the scaling relationships between the L and D and between Apit and D were measured. The scaling exponents between L and D were 1.47, 3.19 and 2.86 for A. rufinerve, M. australis and V. coignetiae, respectively, whereas those between Apit and D were 0.242, 2.11 and 2.68, respectively. Unlike the inter-specific relationships, the wall resistivity fraction (Rwall/(Rlumen + Rwall)) within xylem changed depending on D. In M. australis and V. coignetiae, this fraction decreased with increasing D, while in A. rufinerve, it increased with D. Vessels with a high wall resistivity fraction have high Rwall and total resistivity but are expected to have low susceptibility to xylem cavitation due to a small cumulative Apit. In contrast, vessels with a low wall resistivity fraction have low Rwall and total resistivity but high susceptibility to xylem cavitation. Because the wall resistivity fraction varies with D, the stem xylem contains vessels with different hydraulic efficiencies and safety to xylem cavitation. These features produce differences in the hydraulic properties of plants with different life forms.