Effects of Rootstocks and Interstocks on the Xylem Sap Composition in Apple Trees: Effects on Nitrogen, Phosphorus, and Potassium Content

This paper records the concentration of N, P, and K in sap whichexuded from the cut stumps of composite orchard apple trees.The trees were either single-worked on different rootstocksor incorporated various interstocks. While the volume of sap exuded per unit area of cut surfacedid not appear to vary with rootstock or interstock, exudatefrom the dwarfing rootstocks or interstocks contained depletedlevels of nutrients. A possible mechanism for rootstock and interstock effects isdiscussed in the light of these results.     

苹果树蒸腾规律及其与冠层微气象要素的关系

采用由热扩散植物液流技术测算得到的、周期为2 0 0 3年1～12月、时间步长10 min的苹果树蒸腾数据,结合同步观测得到果树冠层微气象要素值,分析太行山低山丘陵区10年生苹果树蒸腾耗水规律及其与微气象要素关系,旨在为该地区苹果生产提供必要的水分生态理论依据,并力图进一步完善苹果树蒸腾耗水理论。结果表明:(1)苹果树蒸腾(Tr)具有明显时间变化特征。Tr在晴天-多云日的主要和非主要生长季节日变化都表现为单峰曲线趋势。在阴天日,主要生长季节表现为多峰曲线趋势,非主要生长季节单峰除10月和3月份表现为双峰曲线趋势外,其余各月均为单峰曲线趋势。Tr日际变化或季节变化特征是3～4月份逐渐升高、5～6月份达到高峰值、7月份逐渐降低、10月末及11月初Tr迅速降低。全年Tr值为6 0 0 .9mm ,其中,主要生长期间(4～9月份)、非主要生长期间(1～3月份、10～12月份) TR值分别为5 0 2 .6、98.3mm ,分别占全年的83.6 %、16 .4 %。(2 )主要生长期内,Tr与冠层净辐射(Rn) ,空气温度(Ta)、湿度(RH) ,风速(V)等气象要素有很好的复相关性,统计回归方程为Tr =0 .2 74 0 .0 4 2 V 0 .0 0 6 7Ta - 0 .0 14 RH (n=2 6 35 2 ,Tr为单株蒸腾量,Tr、V、Ta、Rn的单位分别为L/ h、m/ s、℃、w/ m2 ,RH为无量纲值,以%表示) ,R2 =0 .     

Specialized 16SrX phytoplasmas induce diverse morphological and physiological changes in their respective fruit crops

The host-pathogen combinations—Malus domestica (apple)/`Candidatus Phytoplasma mali´, Prunus persica (peach)/`Ca. P. prunorum´ and Pyrus communis (pear)/`Ca. P. pyri´ show different courses of diseases although the phytoplasma strains belong to the same 16SrX group. While infected apple trees can survive for decades, peach and pear trees die within weeks to few years. To this date, neither morphological nor physiological differences caused by phytoplasmas have been studied in these host plants. In this study, phytoplasma-induced morphological changes of the vascular system as well as physiological changes of the phloem sap and leaf phytohormones were analysed and compared with non-infected plants. Unlike peach and pear, infected apple trees showed substantial reductions in leaf and vascular area, affecting phloem mass flow. In contrast, in infected pear mass flow and physicochemical characteristics of phloem sap increased. Additionally, an increased callose deposition was detected in pear and peach leaves but not in apple trees in response to phytoplasma infection. The phytohormone levels in pear were not affected by an infection, while in apple and peach trees concentrations of defence- and stress-related phytohormones were increased. Compared with peach and pear trees, data from apple suggest that the long-lasting morphological adaptations in the vascular system, which likely cause reduced sap flow, triggers the ability of apple trees to survive phytoplasma infection. Some phytohormone-mediated defences might support the tolerance.     

Xylem Sap Composition in Apple Trees: Effect of the graft union

This paper records studies on the xylem sap from above and belowthe graft unions of apple trees with M·9, M·7and MM.104 rootstocks. The dry weight of solutes in sap fromabove the unions of the M·9 trees was between 30 and50 per cent lower than in sap from below, while the volume ofsap exuded was not reduced. The graft unions of the M·7and MM·104 trees had no significant effect on the xylemsap. These results are discussed in relation to the dwarfing effectsof rootstocks and interstocks.     

Sorbitol and other carbohydrates in dormant apple shoots as influenced by controlled temperatures

The influence of controlled temperatures on levels of sorbitol and other carbohydrates was determined to provide further information on dormancy of apple trees (Malus sylvestris Mill.). For 3 years, 2-year-old “Red Delicious” apple shoots were collected from mature trees in an orchard at intervals during the autumn and winter, and shoots were stored for 6 hr to 1 week at temperatures from 18 to ?60 °C. Sorbitol and other carbohydrates were estimated in the sap or wood by gas chromatography.Controlled temperatures had a marked influence on the carbohydrate content of excised 2-year-old apple shoots. Levels of sorbitol in the sap were maximum at ?0.6 °C. The increase was greatest at the earliest sampling before complete hardening had occurred in each of the 3 years tested. Total sorbitol in the wood was less influenced by storage at various controlled temperatures than sorbitol in the sap. Levels of fructose, glucose, and sucrose in the wood were higher at temperatures below ?0.6 °C than at warmer temperatures. Levels of starch were usually inversely related to soluble sugars.     

晋西黄土区苹果树边材液流速率对环境驱动的响应

为明确环境因子对树木蒸腾过程的驱动机制,以晋西黄土残塬沟壑区的苹果树(红富士)为对象,利用热扩散式液流技术监测生长季苹果树树干液流的动态变化,并同步监测了气象和土壤水分等环境要素的季节动态.结果表明: 在众多环境因子中,太阳净辐射(R_n)、大气水分亏缺(VPD)与液流速率(J_s)间的相关性最强.在小时或日尺度,环境因子主成分分析中前3个主成分的累积方差贡献率均在86%以上.其中,第一主成分主要包含VPD、R_n等因子,方差贡献率达52%(小时尺度)和63%(日尺度)以上,可归为蒸发需求因子(EDI),是影响该地区果树树干液流的关键综合环境要素集;第二主成分主要包括土壤含水率(SWC)等因子,归为土壤水热供给因子;第三主成分主要包括风速等因子,归为大气水热散失动力因子.在小时或日尺度上,J_s对两种环境因子综合变量(EDI或潜在蒸发散ET₀)的响应都呈显著的指数增长关系,在小时尺度上,基于EDI模拟苹果J_s的指数模型精度更高(R²=0.72),在日尺度上,基于ET₀模拟苹果J_s的指数模型模拟精度更高(R²=0.88).研究结果对于明确苹果树水分传输对环境驱动的响应规律,根据气象要素估算苹果树蒸腾耗水量,并指导果园水分管理均具有重要意义.     

The relationship between nutrients in extracted xylem sap and the susceptibility of fruit trees to silver-leaf disease caused by Stereum purpureum (Pers.) Fr.

Xylem sap was extracted from cut branches of stone-fruit and pip-fruit trees at different times of the year, and analysed for nitrogen and carbohydrate content. Growth of Stereum purpureum was measured in the different sap samples in vitro. Nitrogen and carbohydrate levels in xylem sap from peach trees were highest in late winter and early spring, and these samples supported most growth of S. purpureum. At full bloom the levels of nitrogen and carbohydrate in xylem sap from stone-fruits (peach, nectarine, plum, cherry) were several times greater than in sap from pip-fruits (apple, pear), and the growth of S. purpureum in sap from stone-fruits was approximately ten times that in sap from pip-fruits. The results suggest that seasonal variations in susceptibility of stone-fruits to silver-leaf disease, and the greater susceptibility of stone-fruits than of pip-fruits, are due to differential ability of their xylem saps to support growth of S. purpureum.     

Sorbitol in Tracheal Sap of Apple as Related to Temperature

The influence of dormant-season temperatures on levels of sorbitol was determined in an effort to provide further information on the possible role of sorbitol in dormancy of apple tress (Malus sylvestris Mill.). Two-year-old shoots were collected throughout the dormant season; sorbitol and sugars were determined in tracheal sap extracted under vacuum and in ground dried wood extracted in a soxhlet apparatus. Sorbitol and sugar trimethylsilyl derivatives were detected by gas chromatography. Levels of sorbitol in the sap generally increased during subfreezing temperatures and decreased during warm periods throughout the dormant season. Early peak increases in sap sorbitol appeared to coincide with the beginning of leaf senescence. Postharvest levels of reducing sugars, sorbitol, and particularly sucrose in the wood, increased as the temperature decreased during the dormant season, and the sugar levels decreased with warming temperatures in the spring. Our data indicate that sorbitol and sucrose are important reserves of storage carbohydrates in resting apple trees.     

Transpiration measurements on apple trees with an improved stem heat balance method

Since the late eighties a handy and user-friendly sap flow meter (Dynagage®) is on the market which can quantify 0205 the sap flow through intact plant stems, based on the stem heat balance method. The documentation about its accuracy and reliability, however, is still too limited to use it as a standard method in field experiments with apple trees. We therefore tested this commercial system on potted apple trees (Malus domestica L.; cv. Red Elstar and Jonagold; on rootstock M9 vf) with stem diameters of 1.8 to 4 cm. The measured sap flow was compared with mass loss measured by an automated balance, supposing the total mass loss of the trees was equal to the water loss by transpiration. The results revealed three major problems:

Effects of branch solid Fe sulphate implants on xylem sap composition in field-grown peach and pear: changes in Fe, organic anions and pH

The effects of placing solid implants containing Fe sulfate in branches of Fe-deficient pear and peach trees on the composition of the xylem sap have been studied. Iron sulfate implants are commercially used in northeastern Spain to control iron chlorosis in fruit trees. Implants increased Fe concentrations and decreased organic acid concentrations in the xylem sap, whereas xylem sap pH was only moderately changed. The citrate to Fe ratios decreased markedly after implants, therefore improving the possibility that Fe could be reduced by the leaf plasma membrane enzyme reductase, known to be inhibited by high citrate/Fe ratios. In peach, the effects of the implants could be observed many months post treatment. In pear, some effects were still observed one year after the implants had taken place. Results obtained indicate that solid Fe sulfate implants were capable of significantly changing the chemical composition of the xylem sap in fruit trees.     

Sap feeding by the marsupial Petaurus australis: an enigmatic behaviour?

Summary The Yellow-bellied Glider, Petaurus australis, was observed to feed sporadically but extensively on eucalypt sap. Gliders extracted sap by making incisions into the bark with their procumbent lower incisor teeth and licking the resulting exudate. Less than 1% of possible trees were incised for sap. Sap flow and sap-sugar concentration were estimated for eight sap-site trees and eight non sap-site trees at intervals during an 18 month period. Measurements of sap-sugar concentration differed substantially among trees but fluctuated between sample periods independent of glider use. Sap flow measurements, were always higher for those trees currently in use by gliders, although the particular sap-site trees comprising this group varied between sample periods. The variation in sap flow occurred indenpendent of the incidence of rainfall or the type of microhabitat the tree occupied. Experimental incising of trees, designed to mimic the effects of feeding by gliders, failed to show any effect on sap flow. This suggests that the incidence of sap feeding is determined by a tree's pattern of sap flow and that there may be trees with unusual patterns of sap flow which gliders select as the most favourable trees to incise.     

Studies on a virus causing stem grooving and graft-union abnormalities in Virginia Crab apple*

A virus was transmitted from apple trees to Nicotiana glutinosa and Chenopodium spp. and back to a range of woody indicators in which it affected only Virginia Crab; symptoms were grooves in the xylem, and swelling and necrosis of the scion immediately above the union with the stock. The virus was distinct from that causing stem pitting in Virginia Crab, because although easily detectable in several apple varieties, it was not found in many trees infected with stem pitting virus. The stem grooving virus has flexous particles 600–700 m/μ long, a heat inactivation point of 67 °C, a dilution end-point of 10^-3 in N. glutinosa sap and remains infective for at least 2 days at 20 °C.     

Whole-tree sap flow is substantially diminished by leaf herbivory

Ecohydrological models consider the relationship between tree size and structure (especially leaf area index) and water use but generally treat herbivory as a source of unwanted noise in the data. Little is known of how insect damage to leaves influences whole-plant water use in trees. Water use is driven by environmental demand and the total leaf area through which transpiration can occur, but the effects of insects are expected to be complex. Different kinds of insects could have different effects; for example, chewing insects reduce leaf area, whereas sucking and tissue mining insects reduce leaf function without reducing area. Further, plants respond to herbivory in a range of ways, such as by altering leaf production or abscising leaves. We examined the effect of insects on Eucalyptus blakelyi in a woodland near Canberra, Australia, using sap flow velocity as a measure of whole-plant water use. We applied insecticide to 16 trees matched to an untreated control group. After 6 months, we examined the effects on sap flow velocity and crown condition. There was a general increase in sap flow velocity as trees produced leaves over the growing season, but the increase in sap flow for trees without insecticide protection was half that of the protected trees (increase: 4.4 vs. 9.0 cm/h, respectively). This dramatic effect on sap flow was consistent with effects on crown condition. Unprotected trees had 20% less leaf mass per unit stem in the crown. In addition, unprotected trees had a 20% greater loss of leaf functional area from necrosis. It should be noted that these effects were detected in a year in which there was not an outbreak of the psyllids (Homoptera) that commonly cause severe leaf damage to this tree species. It is predicted that the effect in a psyllid outbreak year would be even more substantial. This result underscores the significant impact that insect herbivores can have on an ecological process of significance to the ecosystem, namely, the movement of water from the soil to the atmosphere.     

An evaluation of hypotheses to explain the pattern of sap feeding by the yellow-bellied glider,Petaurus australis

Yellow-bellied gliders on occasion extensively incise the trunks of eucalypts to feed on sap but in doing so show strong preference for the species and number of trees utilized. Three hypotheses that may account for aspects of this behaviour were examined. They were: (i) sap feeding is related to the incidence of rain; (ii) sap feeding is influenced by the sap flow idiosyncracies of trees; and (iii) sap feeding is influenced by the availability of alternative food resources. Sap feeding at two sites occurred independent of ‘wet’ or ‘dry’ nights. Sap flow (measured as an index) in two species of eucalypt was highly variable among trees and among sample periods. This is the only hypothesis that accounts for the selectivity of the trees incised. Sap feeding at one site coincided with periods of high sap flow. However, at another site, sap feeding did not occur on measured trees despite their often high sap flow. At such times, flowering trees were abundant and gliders fed extensively on nectar. Thus, elevated sap flow may provide the necessary precondition conducive for sap feeding but may be ignored when other food types are available.     

陇东旱地果园覆沙对苹果树蒸腾耗水及果实品质的影响

针对甘肃陇东旱地苹果园季节性干旱问题,以15年生长富2号苹果树为试材,测定果园覆沙后土壤水分、温度,以及果树主干液流速率、叶片气孔导度、果实品质等指标.结果表明：果园覆盖5 cm厚河沙,2-4月地温升高幅度低于1 ℃,6、7月晴天地温升高2.44 ℃,阴天地温升高2.61 ℃;在果树生长季节土壤含水量始终保持在田间持水量的60%以上.土壤含水量较高的时期（H期）晴天,树干液流曲线呈“几”字形宽峰曲线,覆沙处理液流启动时间较对照提前0.6 h,峰值较对照高25.5%,阴天峰值较对照高165.6%,且液流活动时间延长;土壤含水量较低的时期（L期）晴天,覆沙处理液流呈单峰曲线,启动时间较对照提前0.5～1 h,峰值在794 g·h^-1左右,而阴天液流启动时间较对照提前近1 h,峰值较对照高311.0%.3-7月对照的棵间蒸发量（E_s）较覆沙处理高156.0%,过多的地面蒸发是造成果树缺水的主要原因.覆沙后果实单果质量显著提高,果实硬度略有降低,果实可溶性固形物、维生素C、总糖、有机酸含量均有所提高.     

Whole tree xylem sap flow responses to multiple environmental variables in a wet tropical forest

In order to quantify and characterize the variance in rainforest tree physiology, whole tree sap flow responses to local environmental conditions were investigated in 10 species of trees with diverse traits at La Selva Biological Station, Costa Rica. A simple model was developed to predict tree sap flow responses to a synthetic environmental variable generated by a principle components analysis. The best fit was obtained with a sigmoid function which explained between 74 and 93% of the variation in sap flux of individual trees. Sap flow reached an asymptote where higher light and evaporative demand did not cause sap flux to increase further. Soil moisture had little influence on sap flux. The morphological characteristics of the trees significantly affected sap flow; taller trees responded to changes in environmental variables sooner than shorter trees and high liana cover buffered tree sap flow responses to weather. The effect of species‐specific differences on the model was small; the mean effectiveness of the model was reduced by 6% when parameters were estimated from a single pool of measurements taken from all individuals. The results indicate that sap flow response could be effectively estimated using a simple general model and composite environmental index for these 10 diverse tree species.     

Indole-3-acetic acid level in wood,bark and cambial sap of apple rootstocks differing in growth vigour

Free and conjugated IAA levels were determined in wood, bark and cambial sap of M.9, M.26 and MM.106 apple rootstock genotypes differing in growth vigour. The measurements were done on May 15^th, June 15^th and July 15^th. The level of free IAA in bark and wood of the tested trees varied from 27.0 to 52.7 ng·g⁻¹ f.w. while the conjugated hormone content averaged 3–5 times higher. In the bark and wood samples, the differences in auxin content between rootstock genotypes and the time of stem harvesting were insignificant. The level of free IAA in cambial sap was on average 10 to 20 times higher than in both bark and wood tissues, while the conjugated hormone level varied from none (below detection limit) to 37 ng·g⁻¹ f.w. Content of free IAA level in cambial sap from dwarf M.9 rootstock was significantly lower than that in either of the more vigorous genotypes. In both vigorous rootstocks IAA level in cambial sap remained at a similar level at all sampling dates but M.9 cambial sap showed a trend towards decreasing auxin content later in the growing season.     

基于稳定同位素和热扩散技术的张北杨树水分关系差异

利用稳定氢同位素和热扩散技术研究张北防护林杨树的水分来源和蒸腾耗水,分析确定未退化与退化杨树的水分关系差异.结果表明:在生长季节中退化杨树主要利用0～30 cm土壤水分,未退化杨树主要利用30～80 cm土壤水分,两者的水分来源不同.旱季时,未退化杨树利用深层土壤水分和地下水的比例明显高于退化杨树.雨季中,杨树对0～30 cm土壤水分的利用比例增加,退化杨树增加幅度明显高于未退化杨树,对30～180 cm土壤水分的利用比例均减少.未退化杨树的液流速率大于退化杨树,不同天气中液流速率表现出相似的变化趋势,但未退化杨树液流的启动时间比退化杨树早.相关分析表明,未退化和退化杨树液流速率与土壤温度、风速、太阳辐射、相对湿度、空气温度均呈极显著的相关关系.退化杨树液流速率与土壤温度和空气相对湿度呈极显著负相关,与其他因素呈显著正相关,而未退化杨树仅与空气相对湿度呈极显著负相关,与其他因素均呈显著正相关关系,表明退化和未退化杨树蒸腾耗水易受环境条件的影响.退化杨树液流日累计量明显小于未退化杨树,表明其蒸腾耗水量较少;退化杨树水分来源浅,蒸腾耗水的减少并不能阻止林分退化.     

Differences between fruit-bearing and non-bearing apple spurs in activity of an enzyme system decomposing phloridzin

The activity of an enzyme system decomposing phloridzin was investigated in fruitbearing and nonbearing spurs of apple trees, Landsberger Reinette cv., throughout vegetation. Acetone powder obtained from xylem sap of apple spurs was incubated with phloridzinsubstratum in citric buffer at pH 5.5 for 12, 18 and 24 h at 30 °C. A paper and thin-layer chromatography as well aa a spectrophotometric assay were employed for tentative identification of enzymic degradation products. Phloretic acid (PA), co-factor of IAA-oxidase, as well as phloretin (Pin), and phloroglucinol (PI) were found after the digestion of phloridzin. The chromatographed enzyme reaction products were measured densitometrically. The activity of the enzyme system was estimated by its efficiency in PA production and phloridzin disappearance. Obtained values, expressed in percentages, showed that the enzyme activity in fruitbearing spurs was much higher than in nonbearing ones; 30 and 10% of released PA in July, respectively. Because fruitbearing spurs of the apple tree are possibly additionally supplied with auxin translocated from developing seeds, an adaptive character of the enzyme system producing PA, a known auxin repressor, is suggested.