Water in the stems of sessile oak (Quercus petraea) assessed by computer tomography with concurrent measurements of sap velocity and ultrasound emission

Concurrent measurements of heat pulse velocity and ultrasound acoustic emission were performed on the trunks of adult Quercus petraea plants under different water stress conditions. On the trunk section of the plants the wood density was measured non-destructively using a mobile computer tomograph which measures the attenuation of a collimated beam of radiation which traverses the trunk. By scanning the trunk in different directions, it is possible to map wood density in the section. As wood density is strictly correlated with water content, this method allows evaluation of the water content in the trunk section and the water conditions in the different parts of the section. The computer-tomograph technique is non-invasive and is not influenced by climatic fluctuations. A close agreement was found between wood density and heat pulse velocity; the relationship between these two parameters and ultrasound acoustic emission is discussed. Trunks of sessile oak appear well suited as water storage reservoirs playing an important role in tree survival during extended periods of low soil water availability, especially in the context of global climatic change. Here the computer-tomograph methodology is described and suggestions arc made for further research development.     

华南地区常见树种导管特征与树干液流的关系

为探讨树木结构与功能的关系,对华南地区常见8种树木边材的导管特征进行观察,并利用Granier热扩散探针法测量干、湿季树干的液流密度,分析导管特征与树干液流的关系。结果表明,除红锥(Castanopsis hystrix)有两种导管外,大叶相思(Acacia auriculaeformis)、荷木(Schima superba)、火力楠(Michelia macclurei)、藜蒴(C.fissa)、马占相思(A.mangium)、柠檬桉(Eucalyptus citriodora)、尾巨桉(E.urophylla×E.grandis)的导管类型单一。导管特征在种间存在明显差异,且导管长度、密度和孔径之间存在明显相关性,它们与标准化的边材面积呈现显著相关。湿季液流最大值与导管特征无明显相关性,但整树最大蒸腾速率与导管特征呈显著相关;树木的日蒸腾量与导管特征也有明显相关性。因此,树木的液流速率并不受树干的导管影响;而树干的导管孔径与边材面积间的负相关权衡机制,可以降低树种间由于导管孔径差异引起的树干的水分输送速率的差异性。     

Tree water dynamics non-destructively assessed through sap flow measurements and potential evapotranspiration

Sap flow and potential evapotranspiration rates were analyzed for two coniferous tree species (Douglas-fir and Scots pine) and one broadleaf species (sessile oak) in a mixed Carpineto-Quercetum forest during the growing season 2005. The relationship between sap flow and potential evapotranspiration rates, effective crown area as a measure of the relative transpiration and daily relative proportion of the storage water used for transpiration were used as indicators of the tree water dynamics. These indicators were determined on four consecutive days and all three showed good reliability concerning tree water dynamics.     

Sap flow rate and transpiration dynamics in the full-grown oak (Quercus robus L.) in floodplain forest exposed to seasonal floods as related to potential evapotranspiration and tree dimensions

Sap flow rate and transpiration dynamics were studied in the course of 3 years in a dominant tree species in the floodplain forest,i.e. in the full-grown oak (Quercus robur L.) tree, using the method of trunk heat balance devised by the authors. The investigations were carried out at a period at which regular and marked fluctuation in a relatively high water table usually occurred, culminating in seasonal flooding. High sap flow rate values in the tree were established under conditions of non-limiting water supply in soil (up to 400 kg per day or up to 39 000 kg per vegetation period) and characteristic daily flow curves (rounded with a large amplitude and with the maximum at noon), corresponding to those described theoretically. Relationships were inferred by means of which tree water consumption can be calculated under these conditions on the basis of data measured at meteorological stations. From these equations it follows that the transpiration of the tree canopy amounted to 80% of the potential evapotranspiration. The amount of the used daily tree water reserve was assessed to be 0.4 mm in the seasonal average. The transpiration coefficient reached in climatically distinct years the values of 400 to 700 of the increase in tree dry matter. The area of the so-called effective tree-crown ground plan approximated to the area determined geodetically. The results obtained are useful for both ecophysiological and hydrological studies. Some of the described procedures are convenient for the evaluation of functional tree dimensions and according to them also of the forest stand structure.     

Transpiration and stomatal conductance of mistletoe (<Emphasis Type="Italic">Loranthus europaeus</Emphasis>) and its host plant,downy oak (<Emphasis Type="Italic">Quercus pubescens</Emphasis>)

Sap flow rate was measured in the crown of a solitary specimen of downy oak (Quercus pubescens) infested by mistletoe (Loranthus europaeus). Five oak branches and two mistletoe plants were selected for analysis. The seasonal sum of transpired water expressed per leaf area unit was five times higher in the mistletoe than in the oak. In addition, the diurnal curves of sap flow were different between the plants. In the morning, the sap flow measured in the mistletoe lagged one hour behind the sap flow measured in an oak branch unencumbered by mistletoe. In contrast, no time lag was observed in the evening. The proportion of water transpired at night relative to the total transpiration was 7% in both species. The stomatal conductances derived from the inverted Penman-Monteith equation and their dependence on global radiation and the vapour pressure deficit (D) revealed that D exerts a different behaviour in stomatal control of transpiration in the mistletoe. We also determined that the concentration of calcium in the leaf mass could serve as a proxy for transpiration rate, however the relationship was not proportional.     

Seasonal variations of gas exchange, photosynthetic pigments, and antioxidants in Turkey oak (Quercus cerris L.) and Hungarian oak (Quercus frainetto Ten.) of different age

Seasonal changes in physiological and biochemical parameters were studied in 35-, 55- and 140-year-old trees of Turkey oak (Quercus cerris L.) and Hungarian oak (Q. frainetto Ten.), growing in natural stands in Eastern Balkan Mountains (Bulgaria). During the seasonal drought period (August), assimilation activity, transpiration rate, stomatal conductance and water potential had a seasonal minimum in all the studied tree ages and species. The foliar concentrations of glutathione, ascorbate, α-tocopherol, as well as photosynthetic pigments in oak leaves were significantly affected by season. With the increasing age of the studied trees, we observed a decrease of the physiological activity and an increase of the antioxidants’ accumulation. Both the species were drought tolerant and anisohydric, where Q. frainetto exhibited higher rates of gas exchange than Q. cerris. Moreover, they differed in the extent of increase in the foliar antioxidants and carotenoids.     

油松、栓皮栎树干液流速率比较

应用TDP(ThermalDissipationProbe)技术对油松和栓皮栎树干液流进行了初步研究,经过野外近1a的实地定位观测,研究结果显示:栓皮栎月平均树干液流速率在整个生长期都较油松的月平均树干液流速率要高。前者大约是后者的5~10倍。栓皮栎在土壤干旱时期能够在白天产生明显的树干液流。在土壤干旱时期油松白天不产生树干液流而在晚上产生明显树干液流。在土壤相对湿润时期,油松和栓皮栎树干液流速率的波形与太阳总辐射的波形变化一致,但不同的是油松的树干液流速率波形呈明显的单峰状,而栓皮栎树干液流速率波形呈明显的多峰状。在土壤相对湿润时期太阳总辐射很低时能对油松树干液流速率产生明显的降低作用,而对栓皮栎树干液流则没有明显影响。在土壤干旱时期,油松和栓皮栎树干液流速率的峰值分别大约为0.0001cm/s和0.0006cm/s左右;在土壤水分充足时期,油松和栓皮栎树干液流速率的峰值分别大约相等约为0.0015cm/s左右,分别是油松和栓皮栎在干旱日期的液流速率峰值的10倍和2.5倍。     

Seasonal variation of water uptake of a Quercus suber tree in Central Portugal

Hydraulic redistribution (HR) is the phenomenon where plant roots transfer water between soil horizons of different water potential. When dry soil is a stronger sink for water loss from the plant than transpiration, water absorbed by roots in wetter soil horizons is transferred toward, and exuded into dry soil via flow reversals through the roots. Reverse flow is a good marker of HR and can serve as a useful tool to study it over the long-term. Seasonal variation of water uptake of a Quercus suber tree was studied from late winter through autumn 2003 at Rio Frio near Lisbon, Portugal. Sap flow was measured in five small shallow roots (diameter of 3–4 cm), 1 to 2 m from the tree trunk and in four azimuths and at different xylem depths at the trunk base, using the heat field deformation method (HFD). The pattern of sap flow differed among lateral roots as soil dried with constant positive flow in three roots and reverse flow in two other roots during the night when transpiration ceased. Rain modified the pattern of flow in these two roots by eliminating reverse flow and substantially increasing water uptake for transpiration during the day. The increase in water uptake in three other roots following rain was not so substantial. In addition, the flux in individual roots was correlated to different degrees with the flux at different radial depths and azimuthal directions in trunk xylem. The flow in outer trunk xylem seemed to be mostly consistent with water movement from surface soil horizons, whereas deep roots seemed to supply water to the whole cross-section of sapwood. When water flow substantially decreased in shallow lateral roots and the outer stem xylem during drought, water flow in the inner sapwood was maintained, presumably due to its direct connection to deep roots. Results also suggest the importance of the sap flow sensor placement, in relation to sinker roots, as to whether lateral roots might be found to exhibit reverse flow during drought. This study is consistent with the dimorphic rooting habit of Quercus suber trees in which deep roots access groundwater to supply superficial roots and the whole tree, when shallow soil layers were dry.     

Evidence of hydraulic lift in a young beech and oak mixed forest using 18O soil water labelling

Hydraulic lift (HL) by tree roots in a young, broad-leaved, mixed temperate European forest was investigated during the 2008 growing season by injecting ¹⁸O-enriched soil water at a depth of 75–90 cm under drought conditions experimentally imposed in a rain-exclusion system. Based on sap flow, leaf water potential, 2-D root distribution measurements, soil isotope profiles, and xylem water isotope composition, water acquisition and use by two tree species, beech (Fagus sylvatica) and oak (Quercus petraea) was compared. We showed that, unlike oak, beech experienced a marked decrease in sap flow and predawn leaf water potential with increasing soil drought. This behaviour was logical considering the shallower root system in beech than in oak. Six days after ¹⁸O-labelling, we observed isotopic enrichment in the shallower soil layers. Since the intermediate soil layers did not display any enrichment, our results clearly pointed to hydraulic lift by tree roots. The superficial enrichment that was observed in the vicinity of oak trunks and the increase in the isotopic signature of xylem sap in the oak trees but not in the beech trees confirmed the predominant role of oak in the hydraulic lift at our site. Even though facilitation for water acquisition among species was not observed here, our results suggest a potential positive contribution of species like oak toward maintaining species diversity in mixed forest ecosystems submitted to severe drought events.     

Carbon isotope discrimination and wood anatomy variations in mixed stands of Quercus robur and Quercus petraea

The two most common oak species in western Europe, Quercus robur and Quercus petraea, display different ecological behaviours, particularly with respect to their responses to drought. The ecophysiological basis of this niche difference is not understood well. Here we test the hypothesis that these two species present distinct water use efficiencies (WUEs), using the carbon isotope discrimination approach. Leaves and 13 dated ring sequences were sampled in 10 pairs of adult trees growing side by side. Carbon isotope composition was measured on cellulose extracts. In addition, relationships between carbon isotope discrimination and wood anatomy were assessed at the tree level. Quercus robur displayed a 1·0‰ larger isotopic discrimination than Q. petraea, and therefore a lower intrinsic WUE (?13%). This interspecific difference of isotopic discrimination was quite stable with time and independent of tree radial growth and climate fluctuations. A strong positive correlation was observed between average tree values of earlywood vessel surface area and ¹³C isotopic discrimination. This correlation was even higher with ¹³C of the 1976 dry year (r = 0·86). These observations led to the hypothesis that hydraulic properties of xylem could exert a constraint on leaf gas exchange, resulting in a larger WUE for individuals with smaller vessel cross‐section area.     

Quercus ilex Transpiration as Affected by a Prolonged Drought Period

The effect of an extended drought (from 1992 to 1995) on water relations was assessed on evergreen oak (Quercus ilex L.) in a dehesa ecosystem (Seville, Southern Spain). Diurnal and seasonal transpiration patterns were analysed at leaf (porometry) and whole-tree level (sap flow), focusing on the relationship between tree transpiration rates (E_t) and potential evapotranspiration rates (PET). Daily maximum E_t varied over the year, becoming higher between May and August, and lower between November and April. Annual E_t (169 – 205 mm y^–1) accounted for less than 40 % of annual rainfall. The prolonged drought did not affect the water relations of the Q. ilex, mainly due to strong stomatal regulation avoiding the loss of water. Stomatal control was found in all seasons, although it was stronger in summer. This behaviour leads to low water consumption and low E_t/PET ratios throughout the year (0.05 to 0.27).     

A set of primers for the amplification of chloroplast microsatellites in Quercus

The increase in demand for the certification of oak seed lots, as well as control of the geographical origin of oak wood, has led us to develop powerful genetic markers permitting us to discriminate among provenance regions. With the aim of detecting new chloroplast variants, we have identified 17 potential cpSSRs motifs from available oak sequences and tested their variability among French oak populations. Six loci were polymorphic at the intraspecific level in Quercus petraea and Q. robur. Moreover, conservation of the primer pairs was checked on a set of 21 forest tree species and they were all shown to work well on several Quercus species, and even within Fagacaea.     

Radial patterns of xylem sap flow in non-, diffuse- and ring-porous tree species

We investigated radial patterns of sap flux density and wood properties in the sapwood of young loblolly pine (Finns taeda L.), mature white oak (Quercus alba L.) and sweetgum (Liquidambar styraciflua L.), which represent three major classes of wood anatomy: non-porous (coniferous), ring-porous and diffuse-porous. Radial measurements of xylem sap flux density were made in sections of xylem extending to 20 mm and 20–40 mm from the cambium. These measurements were compared with measurements of the relative water content (R_s) and sapwood specific gravity (ρ_r) of corresponding radial sections. In both hardwood species, sap flow differences were rarely significant between the two depth intervals. In pine, a 59% reduction in daily sap flux density from outer to inner sapwood was found. This could not be accounted for by a 3% drop in R_s; rather, an accompanying 9% reduction in ρ_r indicated a transition between the depth intervals from mature to juvenile sapwood, and is the probable cause of the lower flux rate in the inner xylem of pine.     

林木耗水调控机理研究进展

林木的蒸腾耗水量是造林设计与环境水分研究的重要参数。本文就林木耗水的气孔与非气孔调节机制、木质部空穴和栓塞的发生和恢复机理、树体组织水容等方面进行了综述,对它们在树木水分传输过程中的调控作用和意义开展了探讨。目前在蒸腾气孔调节方面,包括,蒸腾午休、夜间蒸腾、气孔振荡和补偿现象等气孔行为的研究工作有待深入。栓塞木质部和空穴化导管恢复的临界条件与重新充注对植物水分运输的重要生理作用要进一步加强。树体组织水容对树木水分传输和耗水的调控机制问题应加以重视。     

Thermal dissipation probe measurements of sap flow in the xylem of trees documenting dynamic relations to variable transpiration given by instantaneous weather changes and the activities of a mistletoe xylem parasite

The thermal dissipation probe was described in the early 1930s for the demonstration of a volume and mass flow of sap in the conductive elements of the xylem in trees. It was subsequently developed further and is now widely used in physiological ecology including measurements in the field. Thermal dissipation demonstrates the occurrence of sap flow and allows determination of its velocity. Here we report simultaneous continuous measurements of sap flow using the thermal dissipation technique and of transpiration by infrared gas analysis for diurnal and annual cycles in a deciduous and an evergreen oak tree, Quercus robur L. and Quercus turneri Willd., respectively, in a deciduous and an evergreen conifer, Larix decidua Mill. and Pinus griffithii McClell., respectively, and the host/mistletoe consortium of the deciduous linden Tilia mandschurica Rupr. & Max. and the evergreen Viscum album L. We show (1) that in diurnal cycles sap flow closely follows dynamic changes of the rate of transpiration elicited by daily fluctuations of weather parameters (sunshine, cloudiness, air temperature and humidity), (2) that in annual cycles sap flow reflects autumnal yellowing and shedding of leaves of the deciduous trees. We report for the first time comparative measurements of sap flow towards mistletoe shoots and host branches in a parasite/host consortium. This demonstrates (3) that mistletoes maintain considerably larger sap flow rates in their xylem conduits than the adjacent host branches dragging the transpiration stream of their host towards their own shoots. We also show (4) that even after the deciduous host has shed its leaves and itself does not transpire any more the evergreen mistletoe towards its shoots can maintain the persistence of a continuous sap flow via the stem and branches of the host as long as frost does not prevent that. The work presented underlines the contention that transpiration is the driving force for sap flow with continuous files of water in the xylem. It shows for the first time that mistletoes direct the flow of water through host roots and stems towards its own shoots by not only performing stronger transpiration as it is known from the literature but also by maintaining larger sap flow rates in the xylem conduits of its stems.     

Epicormic ontogeny in Quercus petraea constrains the highly plausible control of epicormic sprouting by water and carbohydrates

Background and Aims

There is increasing evidence that suppressed bud burst and thus epicormic shoot emergence (sprouting) are controlled by water–carbohydrate supplies to entire trees and buds. This direct evidence is still lacking for oak. In other respects, recent studies focused on sessile oak, Quercus petraea, have confirmed the important constraints of sprouting by epicormic ontogeny. The main objective of this paper was thus to provide provisional confirmation of the water–carbohydrate control and direct evidence of the ontogenic constraints by bringing together results already published in separate studies on water status and distribution of carbohydrates, and on accompanying vegetation and epicormics, which also quantify epicormic ontogeny.

Methods

This paper analyses results gained from a sessile oak experiment in which part of the site was free from fairly tall, dense accompanying vegetation. This experiment was initially focused on stand water status and more recently on the carbohydrate distribution of dominant trees. External observations of the epicormic composition and internal observations with X-ray computer tomography were undertaken on 60 and six trees, respectively.

Key Results

Sprouting was more intense in the part of the stand free from accompanying vegetation and on upper trunk segments. A clear effect of epicormic ontogeny was demonstrated as well: the more epicormics a trunk segment bears, the more chances it had to bear sprouts.

Conclusions

These results indirectly infer water–carbohydrate control and show direct evidence of constraints by epicormic ontogeny. These results have far-reaching consequences related to the quantification of all functions fulfilled by any type of epicormic structure in any part of the tree.     

黄土丘陵区辽东栎和刺槐树干液流时滞效应与蒸腾特征的关联性

运用Granier热扩散探针法,于2016年7-9月对半干旱黄土丘陵区天然次生林树种辽东栎和人工林树种刺槐的树干液流进行连续测定,并同步监测气象因子和土壤含水量,用错位相关法分析液流通量密度与空气水汽压亏缺日变化的时滞长度,研究2个树种不同径级个体在不同土壤水分条件下液流通量密度与蒸腾驱动因子之间的时滞效应.结果表明:辽东栎和刺槐液流通量密度的日变化节律与气象因子显著相关,空气水汽压亏缺峰值的出现较辽东栎树干液流通量密度滞后118.2 min,较刺槐树干液流通量密度滞后39.5 min;而光合有效辐射的峰值通常滞后于辽东栎12.4 min,提前于刺槐68.5 min.液流通量密度和空气水汽压亏缺的时滞长度与树种和土壤含水量显著相关,辽东栎、刺槐在土壤含水量较高时段的时滞长度分别大于土壤含水量较低时段32.2和68.2 min.时滞长度与径级的相关性整体上未达到显著水平,但在土壤含水量较低时段小径级刺槐的时滞长度大于大径级21.4 min,差异达到了显著水平.两树种液流通量密度与空气水汽压亏缺之间的时滞效应反映了对蒸腾驱动因子的敏感性,较好的土壤水分条件有利于液流通量密度提早达到峰值,较低土壤水分会导致树干液流对气象环境因子响应的敏感性降低;刺槐树干液流受土壤水分的影响更显著.     

太行山南麓低山丘陵区栓皮栎直径变化及其与环境因子的关系

使用周长传感器(Circumference DC2)研究了太行山南麓低山丘陵上栓皮栎人工林树干直径的日变化及其影响因子.结果表明:在季节性干旱期间,栓皮栎树干的直径变化周期性明显,直径收缩与液流启动时间基本一致,直径最小值滞后于液流速率最大值3 ～4 h;栓皮栎直径日最大收缩量(MDS)呈现低-高-低的变化趋势,与累积液流通量和叶片水势日差值极显著相关,与土壤含水量呈显著的二次方程关系.MDS值受气象因子变化的影响,与温度日差值、蒸汽压亏缺日差值和相对湿度日差值显著相关,而与太阳辐射日差值的相关性不显著.连续降水后,土壤水分不再是栓皮栎直径变化的限制因子,MDS值与累积液流通量、叶片水势、土壤含水量和气象因子差值的相关性均不显著.季节性干旱期和雨季的土壤含水量和温度是影响树干直径日变化的主要因子.     

六盘山辽东栎、少脉椴天然次生林夏季蒸散研究

2004年8～9月份,利用热扩散技术,结合微型蒸渗仪和水文学方法,研究了辽东栎、少脉椴次生林蒸散组成及其与林分结构的关系.结果表明,辽东栎和少脉椴树干的液流密度在"相对静止期"内比较稳定和微弱,其值在0.05μl·cm^-2·min^-1以下;在"活跃期"内树干液流密度上升较快,并呈单峰、双峰或多峰曲线,其值在0.25μl·cm^-2·min^-1以下;两树种单株蒸腾量有明显的种间差异,前者晴天和阴雨天单株蒸腾量分别为5.31和2.48 L·d^-1,为后者的2.3倍和3.75倍.林下灰_栒子和黄刺玫蒸腾速率日均值接近,分别为0.331和0.321 g·g^-1·h^-1.次生林日均蒸散量1.4 mm·d^-1,其中蒸腾量0.72 mm·d^-1、土壤蒸发量0.19 mm·d^-1、林冠截留量0.4 mm·d^-1,各占总量的49.6%、13.3%和37.1%.乔、灌木树种组成对次生林蒸腾量影响的表现不同,前者表现为个体蒸腾量的种间差异,而后者取决于单位林地面积上各树种的叶量.乔木层、灌木层和草本层(含土壤层)日均蒸散量分别为0.96、0.30和0.19 mm·d^-1,各占总量的65.8%、20.9%和13.3%,说明乔木层对林分日蒸散量大小起主要作用,灌木层次之,草本和土壤蒸发量的贡献最小.     

Detection and features of wetwood in Quercus mongolica var. grosseserrata

To elucidate the location of water in wetwood, variations in moisture content within a tree of Quercus mongolica var. grosseserrata were observed macroscopically by determination of moisture content and by soft x-ray photography. Then the typical wetwood, which had been precisely located by such macroscopic analysis, was examined further by cryo-scanning electron microscopy. The moisture content varied considerably within the tree trunk, and typical wetwood was detected in limited regions of the heartwood. The distribution of the wetwood did not always correspond to the discoloration of the wood. In the wetwood, almost all the lumina of vessels and fibers were filled with water, while the lumina of ray and axial parenchyma cells were often cavitated. Intercellular spaces produced by deformation of cell walls and radial shakes were filled with water. In addition, tyloses were collapsed and torn in the wetwood, and they did not block the vessel lumina. These results suggest the possibility that vessels and fibers might serve as effective pathways for the accumulation of water in the wetwood.