Tree water transport mediating the changing environmental conditions to tree physiological processes

Teemu Paljakka
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Abstract

Tree vascular tissues connect resource availability to tree physiological processes and growth. The xylem transports water from the soil up to the canopy of even 100-metre tall trees, whereas phloem transport connects the photosynthesis in leaves and the tree metabolic processes, including growth and tree defences against insect and pathogen attacks. Water deficit results in the closing of leaf stomata and decreasing photosynthetic production, as water and carbon dioxide are exchanged through the stomata between the leaf and ambient air. Phloem transport is driven by turgor pressure gradients generated by the interplay of phloem osmotic concentration and xylem water potential. Trees have adapted to local environmental conditions and they adjust to fast environmental changes with physiological responses. This thesis investigates tree physiological responses in vascular tissues, such as osmolality, water potential and stomatal conductance, to environmental conditions in two conifers: Scots pine and Norway spruce. Seasonality in soil temperature and soil water content affect soil-to-leaf hydraulic conductance, and stomatal conductance is connected to these seasonal patterns in water transport. Soil environment is thus mediated to tree functionality through tree water transport. This thesis also supports Münch’s theory that it is plausible to explain phloem transport in conifers in field conditions with osmotic gradients and gravity. Xylem water potential reflects to osmotic potential and turgor pressure of the inner bark by modifying tissue solute and water content. The turgor gradients hence seem to determine daily and seasonal carbon allocation patterns according to water availability. Pathogenic infections may introduce more rapid changes in tree hydraulic conductance through a decrease in xylem sap surface tension and xylem conductivity during massive invasions of bark beetles that vector blue-stain fungi such as Endoconidiophora polonica. These pest attacks weaken tree vitality and may also increase tree vulnerability to hydraulic failure in the xylem.
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树木水分运输介导环境条件变化对树木生理过程的影响
树木维管组织将资源的可利用性与树木的生理过程和生长联系起来。木质部将水分从土壤输送到100米高的树木的树冠,而韧皮部的运输将叶片中的光合作用和树木的代谢过程联系起来,包括生长和树木抵御昆虫和病原体的攻击。水分缺乏导致叶片气孔关闭,光合作用减少,因为水分和二氧化碳通过气孔在叶片和周围空气之间交换。韧皮部运输是由韧皮部渗透浓度和木质部水势相互作用产生的胀压梯度驱动的。树木已经适应了当地的环境条件,它们通过生理反应来适应快速的环境变化。本文研究了两种针叶树:苏格兰松和挪威云杉的维管组织对环境条件的生理反应,如渗透压、水势和气孔导度。土壤温度和含水量的季节变化影响土壤对叶片的水力导度,而气孔导度与水分输送的季节变化有关。因此,土壤环境是通过树木的水分输送来调节树木功能的。这篇论文也支持了m nch的理论,即在渗透梯度和重力条件下解释针叶树韧皮部运输是合理的。木质部水势通过改变组织溶质和含水量来反映内部树皮的渗透势和膨胀压。因此,水位梯度似乎决定了每日和季节的碳分配模式,根据水的可用性。在蓝斑内生真菌(Endoconidiophora polonica)等树皮甲虫大规模入侵期间,病原菌感染可能通过降低木质部汁液表面张力和木质部电导率而引起树木水力电导率的快速变化。这些害虫的攻击削弱了树木的活力,也可能增加树木对木质部水力破坏的脆弱性。
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