Repair of severe winter xylem embolism supports summer water transport and carbon gain in flagged crowns of the subalpine conifer Abies veitchii.

Xylem embolism induced by winter drought is a serious dysfunction in evergreen conifers growing at wind-exposed sites in the mountains. Some coniferous species can recover from winter embolism. The aim of this study was to determine whether wind direction influences embolism formation and/or repair dynamics on short windward and long leeward branches of asymmetrical `flagged' crowns. We analyzed the effect of branch orientation on percentage loss of xylem conductive area (PLC), leaf functional traits and the xylem:leaf area ratio for subalpine, wind-exposed flagged-crown Abies veitchii trees in the northern Yatsugatake Mountains of central Japan. In late winter, the shoot water potential was below -2.5 MPa, and the PLC exceeded 80% in 2-year-old branches, independent of branch orientation within a flagged crown. Both of these parameters almost fully recovered by summer. At branch internodes 4 years of age and older, seasonal changes in PLC were not found in either windward or leeward branches, but the PLC was higher in less leafy windward branches. The leaf nitrogen content and water-use efficiency of mature leaves were comparable between windward branches and leafy leeward branches. The ratio of xylem conductive area to total leaf area was the same for windward and leeward branches. These results indicate that the repair of winter xylem embolism allows leaf physiological functions to be maintained under sufficient leaf water supply, even on winter-wind-exposed branches. This permits substantial photosynthetic carbon gain during the following growing season on both windward and leeward branches. Thus, xylem recovery from winter embolism is a key trait for the survival of harsh winters and to support productivity on the individual level in flagged-crown A. veitchii trees.

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.

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.