Identification and quantitation by GC-MS of zeatin and zeatin riboside in xylem sap from rootstock and scion of grafted apple trees

Zeatin and zeatin riboside were identified by full-scan gas chromatography-mass spectrometry (GC-MS) in xylem sap of clonal apple rootstocks (M.27, M.9 and MM.106). These rootstocks exhibit a wide range of control over tree size when grafted to a common scion. The concentrations of zeatin and zeatin riboside were measured by GC-MS selected ion monitoring (SIM) in shoot xylem sap and root pressure exudate obtained from these rootstocks and from trees of Fiesta scion grafted onto the rootstocks. Zeatin was the predominant cytokinin in xylem sap from the dwarfing rootstocks, M.27 and M.9, while zeatin riboside was the predominant cytokinin in xylem sap from the more invigorating rootstock MM.106. Cytokinin concentrations (ng ml^–1) in root pressure exudate and shoot xylem sap, (i.e. from above the graft union in composite trees), increased with increasing vigour of the rootstock, irrespective of whether the plants were non-grafted rootstocks, or were composite plants of Fiesta scion grafted onto the rootstocks. Cytokinin content (ng shoot^–1) of shoot sap differed with rootstock; the more invigorating (MM.106) had greater amounts of cytokinins than the more dwarfing (M.9 and M.27) rootstocks. These results are discussed in relation to possible influences of roots on the growth of shoots via cytokinin supplies in the xylem sap.     

Influence of rootstock on antioxidant system in leaves and roots of young apple trees in response to drought stress

Grafting rootstocks are widely used to enhance plants resistance to various biologic and abiotic stresses. We determined how the rootstock genotype might influence plant responses to drought, using 2-year-old ‘Gale Gala’ apple trees grafted onto Malus sieversii and M. hupehensis. Under water stress, trees with the former as their rootstock had smaller reductions in rates of relative growth and photosynthesis, total biomass, leaf area, levels of leaf chlorophyll, and relative water content compared with those grafted onto the latter. They also had greater maximum photochemical efficiency and water-use efficiency. On the other hand, trees growing on M. sieversii rootstock had less production of superoxide radicals and hydrogen peroxide in both leaves and roots than those growing on M. hupehensis in response to drought stress. Furthermore, under drought conditions, leaves and roots from trees grafted onto M. sieversii had greater synthesis of ascorbic acid and glutathione, as well as higher activities of superoxide dismutase, catalase, ascorbate peroxidase, monodehydroascorbate reductase, dehydroascorbate reductase, and glutathione reductase. These results suggest that the choice of grafting rootstock can enhance drought resistance by improving the antioxidant system in a plant. Here, ‘Gale Gala’ trees grafted onto M. sieversii were more drought-resistant than those on M. hupehensis rootstock.     

Root and stem hydraulic conductivity as determinants of growth potential in grafted trees of apple (Malus pumila Mill.)

The anatomy of the graft tissue between a rootstock and its shoot (scion) can provide a mechanistic explanation of the way dwarfing Malus rootstocks reduce shoot growth. Considerable xylem tissue disorganization may result in graft tissue having a low hydraulic conductivity (k(h)), relative to the scion stem. The graft may influence the movement of substances in the xylem such as ions, water and plant-growth-regulating hormones. Measurements were made on 3-year-old apple trees with a low-pressure flow system to determine k(h) of root and scion stem sections incorporating the graft tissue. A range of rootstocks was examined, with different abilities of dwarfing; both ungrafted and grafted with the same scion shoot cultivar. The results showed that the hydraulic conductivity (k(hroot)) of roots from dwarfing rootstocks was lower compared with semi-vigorous rootstocks, at least for the size class of root measured (1.5 mm diameter). Scion hydraulic conductivity (k(hs)) was linked to leaf area and also to the rootstock on to which it was grafted, i.e. hydraulic conductivity was greater for the scion stem on the semi-vigorous rootstock. Expressing conductivities relative to xylem cross-sectional areas (k(s)) did not remove these differences suggesting that there were anatomical changes induced by the rootstock. The calculated hydraulic conductivity of the graft tissue was found to be lower for grafted trees on dwarfing rootstocks compared to invigorating rootstocks. These observations are discussed in relation to the mechanism(s) by which rootstock influences shoot growth in grafted trees.     

Changes in response to drought stress of triticale and maize genotypes differing in drought tolerance

Direct effects and after-effects of soil drought for 7 and 14 d were examined on seedling dry matter, leaf water potential (ψ), leaf injury index (LI), and chlorophyll (Chl) content of drought (D) resistant and sensitive triticale and maize genotypes. D caused higher decrease in number of developed leaves and dry matter of shoots and roots in the sensitive genotypes than in the resistant ones. Soil D caused lower decrease of ψ in the triticale than maize leaves. Influence of D on the Chl b content was considerably lower than on the Chl a content. In triticale the most harmful D impact was observed for physiologically younger leaves, in maize for the older ones. A period of 7-d-long recovery was too short for a complete removal of an adverse influence of D.     

Growth, gas exchange, and water-use efficiency response of two young apple cultivars to drought stress in two scion-one rootstock grafting system

The combination of two scion-one rootstock was used for two apple cultivars, ‘Pink Lady’ and ‘Qinguan’, budded on the same, one-year-old Malus hupehensis (Pamp.) Rehd. to reduce the impact of root and pot size and in order to understand the growth, water-use efficiency (WUE), and chlorophyll fluorescence characteristics. The two-scion grafted trees were planted in plastic pots under two water regimes, i.e. 70% field capacity (FC) and 55% FC. Results indicated that different scions were affected differently by drought stress. ‘Pink Lady’ had higher net photosynthetic rate (P _N), stomatal conductance (g _s), and transpiration rate (E) compared with ‘Qinguan’ under both water treatments. However, ‘Qinguan’ had lower minimal fluorescence (F₀), higher maximum fluorescence (F_m), and higher maximum photochemical efficiency of photosystem II (F_v/F_m) than ‘Pink Lady’ at 55% FC. Moreover, ‘Qinguan’ had larger shoot dry mass (ShDM) and higher intrinsic WUE_I than ‘Pink Lady’ under both water status. Gas-exchange and growth parameters, except for P _N and scion diameter, were significantly affected by the cultivar and water treatment. At 70% FC, ShDM was significantly correlated with WUE_I. Moreover, WUE_I was negatively linearly correlated with g _s at either 70 or 55% FC. These results might indicate that ‘Pink Lady’ was more sensitive to drought than ‘Qinguan’. ‘Qinguan’ apple was able to improve WUE more than ‘Pink Lady’ under both well-watered and drought conditions. The growth parameters and photosynthetic capacity of two different scions showed that the combination of double scion-one rootstock might eliminate the influences of the rootstock and pot size.     

基于茎干直径微变化制定苹果灌溉制度

茎干直径的动态微变化是研究植物体水分和生长状况的重要指标。利用测树器监测西北旱区盛果期苹果树茎干直径微变化规律,根据监测记录获得茎干直径日最大值(MXTD)、茎干直径日最大收缩量(MDS)数据,并探讨茎干直径微变化规律及其对环境因素的响应,为茎干直径微变化用于指导精确灌溉提供科学依据。实验结果表明,晴天或多云天气下,苹果树茎干直径在每天的7:00或8:00时刻达到一天最大值,在16:00左右达到一天的最小值,茎干直径年增长量与果实产量成反比例关系。整个生育期MXTD呈先快速增加后平稳的变化特征。2010年MDS与茎干水势(φstem)呈显著负相关关系(r2=0.76***,n=14),这表明MDS可以反映苹果树的水势状况。生育后期的MDS对环境因素响应比生育前期敏感,全生育期MDS与气象因素的决定系数大小顺序为日最大水汽压差(VPDmax)日最高温度(Tmax)净辐射(Rn)。茎干直径微变化规律可以反映西北旱区盛果期果树的水分状况,可以为果园灌溉制度的确定提供科学依据。     

6个耐旱树种木质部结构与栓塞脆弱性的关系

木质部栓塞脆弱性对干旱响应的研究已成为全球气候变化背景下的热点和重要内容。该文以6个耐旱树种刺槐(Robinia pseudoacacia)、沙棘(Hippophae rhamnoides)、榆树(Ulmus pumila)、元宝枫(Acer truncatum)、旱柳(Salix matsudana)、榛(Corylus heterophylla)为研究对象, 采用Cochard Cavitron离心机技术建立木质部栓塞脆弱曲线, 计算木质部栓塞脆弱性, 利用染色法、硅胶注射法等测定木质部导管直径、导管内径跨度、导管连接度、导管密度、导管长度和木质部密度, 探究木质部结构与栓塞脆弱性的关系, 区分6个耐旱树种木质部结构在抗栓塞性上的差异, 以期建立6个耐旱树种在木质部结构方面的抗栓塞性指标。结果表明: 6个耐旱树种木质部栓塞脆弱性大小为刺槐>榆树>沙棘>旱柳>元宝枫>榛, 其中, 刺槐、沙棘和榆树的栓塞脆弱曲线为“r”形, 而元宝枫、旱柳和榛的栓塞脆弱曲线为“s”形, 脆弱曲线为“r”形的树种与脆弱曲线为“s”形的树种栓塞脆弱性差异极显著(p < 0.01)。线性分析表明: 木质部结构影响各树种的栓塞脆弱性, 其中, 木质部密度影响最大(t = 0.702), 导管直径次之(t = 0.532), 导管长度影响最小(t = 0.010)。     

Clarifying the effects of dwarfing rootstock on vegetative and reproductive growth during tree development: a study on apple trees

BACKGROUND AND AIMS: Despite the widespread use of dwarfing rootstocks in the fruit-tree industry, their impact on tree architectural development and possible role in the within-tree balance between growth and flowering are still poorly understood, in particular during the early years of growth. The present study addressed this question in apple trees, through a detailed analysis of shoot populations, i.e. both vegetative and flowering shoots, during tree development. METHODS: Architectural databases were constructed for trees of two cultivars that were either own-rooted or grafted on dwarfing rootstock. Within-tree shoot demographics and annual shoot characteristics, i.e. their dimensions, number of laterals and flowering, were observed from the first to the fifth year of growth and compared among scion/root system combinations. KEY RESULTS: Differences in axis demographics appeared among scion/root system combinations after the second year of growth. Differences were found (a) in the number of long axes and (b) the number of medium axes. Dwarfing rootstock reduced the total number of axes developed in a tree, and this reduction resulted from proportionally more medium axes and spurs than long axes. The life span of spurs was also shortened. These phenomena appeared after an increase in flowering that started in the second year of growth and involved both axillary and terminal positions. Flowering regularity was also increased in grafted trees. CONCLUSIONS: These results confirm that the number of long shoots and flowering potential depend on the cultivar. They indicate that tree architectural plasticity in response to its root system mainly derives from the number of medium shoots developed and follows priorities within the whole tree axis population. There was also evidence for dwarfing rootstock involvement in adjusting the flowering abundance and that differences in flowering occurrence take precedence over those regarding vegetative growth during tree development.     

Cellular boron allocation and pectin composition in two citrus rootstock seedlings differing in boron-deficiency response

Aims

Variation in boron (B) efficiency in citrus in different rootstock genotypes is expressed as large differences in the occurrence of leaf symptoms and dry mass production under low B conditions, but the mechanisms responsible for such differences are unknown. This paper aims to determine whether differences in B uptake, cellular B allocation, and pectin content can explain genotype differences in B efficiency between B-efficient citrange (Citrus sinensis (L.) Osb. × Poncirus trifoliata (L.) Raf.) and B-inefficient trifoliate orange (Poncirus trifoliata (L.) Raf.) citrus rootstock.

Methods

Plants were grown hydroponically in a nutrient solution supplemented with 5 μM B for 14 days and then transferred to a B-free medium (0 μM B) or control medium (5 μM B) for 35 days. Boron uptake and allocation and cell wall pectin contents were examined.

Results

After 35 days under B deprivation, shoot dry mass in trifoliate orange decreased by 28 %, but shoot dry mass of citrange was not significantly affected. Root growth of both types of rootstock seedlings was inhibited, but the trifoliate orange was affected more than the citrange. In comparison with B concentrations in plants prior to the commencement of B treatments, B deprivation for 35 days decreased B concentration in various parts of citrange plants, and the reduction was much greater in trifoliate orange plants. Trifoliate orange seedlings contained higher B concentration and total B in cell wall on a dry leaf basis than citrange subject to 5 μM B treatment. However, the proportion of leaf B allocated in cell wall was higher in citrange than trifoliate orange when B supply was deficient in the nutrient. The changes in pectin composition in cell wall due to B deprivation differed between citrange and trifoliate orange. The decreased uronic acid (UA) content in the Na₂CO₃-soluble pectin was observed in both rootstock, but the increased UA content in CDTA-soluble pectin was observed only in citrange.

Conclusions

These results demonstrated that a combination of greater B uptake ability, greater B accumulation in cell walls, as well as the increased CDTA-soluble pectin, under limited external B supply, contribute to the integrity of cell walls in citrange and therefore increased tolerance to B deficiency.     

Modification of gibberellin biosynthesis in the grafted apple scion allows control of tree height independent of the rootstock

The availability of short stature apple scions that required minimal applications of chemical growth retardants and could be used with a range of rootstocks would be of considerable benefit to fruit growers. We have suppressed the expression of a gene encoding the gibberellin (GA) biosynthetic enzyme GA 20-oxidase to reduce the levels of bioactive GAs in a scion variety, resulting in significant reductions in stem height. Application of GA3 reversed the effect. The scion remained dwarfed after grafting on to normally invigorating rootstocks, whilst control plants of the same cultivar displayed the expected vigour when grafted on to these rootstocks. This approach could be applicable to any perennial crop variety, allowing dwarf trees to be obtained on any available rootstock or on their own roots without the need for chemical growth retardant application. In effect, seedlings that are well suited to local conditions (drought, salinity) could be employed as tree rootstocks, as could existing rootstocks valued for characters other than vigour control, such as pest and disease resistance.     

Effect of rootstock on photoperiodic control of elongation growth in grafted ecotypes of Salix

Scions of a southern (59° 40'N Lat.) and a northern (69° 39'N Lat.) ecotype of Salix pentandra L. were grafted on clonal rootstocks of the same ecotypes. Effects of photoperiod on elongation growth of the 4 combinations (south/south, south/north, north/south and north/north) were studied in a phytotron at 18° and 15°C. The photoperiodic response was primarily dependent on the ecotype of the scion, but this response was significantly modified by the rootstock. Cessation of apical growth was advanced by a northern clone and delayed by a southern clone as a rootstock. The results indicate that the critical photoperiod for cessation of apical growth could be slightly decreased by a northern and increased by a southern rootstock.     

Wood diameter indicates diurnal and long-term patterns of xylem water potential in Norway spruce

The stem diameter of adult Norway spruce trees was measured to see whether changes in xylem water potential lead to detectable radial deformation of the wood. The dendrometers used in these experiments measured only the dimensional changes of the woody cylinder (sap- and heartwood). Wood diameter was measured close to the ground and just below the living crown. After correction for thermal expansion of dendrometers and wood, diurnal variation of wood diameter ranged between 50 and 180 µm. Psychrometric measurements showed that xylem water potential varied in parallel to wood diameter. Diameter changes were always more pronounced at the higher stem position and exhibited a clear diurnal pattern. During the day, wood diameter decreased with increasing vapor pressure deficit and transpiration rate and with decreasing twig water potential. At night, the wood re-expanded but did not always reach the dimension of the previous day. Pre-dawn wood diameter decreased during periods of soil drought, a process which rapidly stopped and reversed after rain events. On several days, oscillation in wood diameter was observed during the mid-day hours. The oscillation had a period of approximately 50 min and showed a phase shift between different stem heights. All observed patterns of wood shrinkage and expansion were consistent with the hypothesis that xylem water tension leads to an elastic contraction of xylem conduits. The results demonstrate that xylem diameter is more suitable than whole-stem diameter for monitoring changes in xylem water potential.     

Effect of fruiting and drought or flooding on carbon balance of apple trees

The response of fruiting or deblossomed trees to water stress such as drought or flooding was investigated in six semi open-top cuvettes each containing one apple (Malus domestica Borkh. cv. Golden Delicious) tree. Xylem water potentials of leaves dropped from -1.2 to -4.1 MPa within 7 d of drought, the effect being enhanced by fruiting. Apple trees without fruits showed smaller reductions in net photosynthetic rate (P _N ) and dark respiration rate (R _D ) after 2 d of drought and hence more positive carbon balances relative to fruiting trees. Flooding for 4 d had a more pronounced effect on P _N than on transpiration, resulting in a reduced water use efficiency (WUE). This reduction in WUE was greater in the non-fruiting trees. Flooding reduced P _N of the whole apple canopies irrespective of fruiting; aple trees without fruits increased R _D resulting in a less positive carbon balance relative to fruiting trees. Fruiting increased the sensitivity to drought of apple trees (R _D and P _N ), but decreased their sensitivity to flooding (R _D and WUE), suggesting different adaptation mechanisms for the two forms of water stress. This revised version was published online in August 2006 with corrections to the Cover Date.     

太行山南麓不同径级栓皮栎生长对气候要素及干旱事件的响应

为评估径级对树木的气候-生长关系的影响,建立太行山南麓低海拔地区栓皮栎全轮、早材、晚材宽度年表,对比两个径级栓皮栎人工林径向生长对气候响应的敏感性差异,使用叠加时代分析揭示干旱事件对不同径级栓皮栎的影响,为气候变化背景下研究区栓皮栎人工林可持续经营提供参考数据.结果 表明:大径级栓皮栎全轮、晚材年表的平均敏感度高于小径...     

A dendroclimatological approach to diameter growth in adult cork-oak trees under production

The diameter growth of mature cork oaks under cork production in southwestern Portugal was studied during a 9-year cork-production cycle in relation to fluctuations in precipitation and temperature, and to drought indices combining the two variables. A dendroclimatological approach was applied to the series of 8 complete years contained in the cork-production cycle and growth effects were removed by standardization and calculation of annual growth indices. The most important factor positively influencing diameter growth was cumulative precipitation in the growing season (January-June) and in the previous autumn and winter (October/November of the previous year-June). The monthly temperature is a less influential factor, negatively correlated with diameter growth in high summer drought (August) and positively correlated at the beginning of physiological activity (March).     

Differences in xylem and leaf hydraulic traits explain differences in drought tolerance among mature Amazon rainforest trees

Considerable uncertainty surrounds the impacts of anthropogenic climate change on the composition and structure of Amazon forests. Building upon results from two large‐scale ecosystem drought experiments in the eastern Brazilian Amazon that observed increases in mortality rates among some tree species but not others, in this study we investigate the physiological traits underpinning these differential demographic responses. Xylem pressure at 50% conductivity (xylem‐P₅₀), leaf turgor loss point (TLP), cellular osmotic potential (π_o), and cellular bulk modulus of elasticity (ε), all traits mechanistically linked to drought tolerance, were measured on upper canopy branches and leaves of mature trees from selected species growing at the two drought experiment sites. Each species was placed a priori into one of four plant functional type (PFT) categories: drought‐tolerant versus drought‐intolerant based on observed mortality rates, and subdivided into early‐ versus late‐successional based on wood density. We tested the hypotheses that the measured traits would be significantly different between the four PFTs and that they would be spatially conserved across the two experimental sites. Xylem‐P₅₀, TLP, and π_o, but not ε, occurred at significantly higher water potentials for the drought‐intolerant PFT compared to the drought‐tolerant PFT; however, there were no significant differences between the early‐ and late‐successional PFTs. These results suggest that these three traits are important for determining drought tolerance, and are largely independent of wood density—a trait commonly associated with successional status. Differences in these physiological traits that occurred between the drought‐tolerant and drought‐intolerant PFTs were conserved between the two research sites, even though they had different soil types and dry‐season lengths. This more detailed understanding of how xylem and leaf hydraulic traits vary between co‐occuring drought‐tolerant and drought‐intolerant tropical tree species promises to facilitate a much‐needed improvement in the representation of plant hydraulics within terrestrial ecosystem and biosphere models, which will enhance our ability to make robust predictions of how future changes in climate will affect tropical forests.     

A cybernetic model of growth correlation in young apple trees

Growth correlations among lateral shoots of one-year-old apple treesMalus Mill., were described. The existence of a mechanism was postulated through which small original differences between buds or young shoots are quickly augmented, thus leading to differentiation of long shoots and short shoots. Existing hypotheses do not seem sufficient for explaining correlative phenomena of this type; a cybernetic approach was therefore applied. Studying growth correlations in terms of cybernetic revealed that previous hypotheses concerning correlations do not contradict each other as often thought, but depict different links in a more general chain of events. A cybernetic model points out the importance of root influences in interaction among shoots. It also shows that synergism between auxin, gibberellin and cytokinin in xylem and phloem differentiation and in starch metabolism is very important for understanding correlations in apple trees.     

Relations between vulnerability to xylem embolism and xylem conduit dimensions in young trees of Quercus corris

Measurements of xylem conduit length and width and the distribution of xylem conduit ends were made in inter-nodes (I), nodes (N) and twig junctions (J) of 1-, 2- and 3-year-old twigs of plants of Quercus cerris L. Parallel measurements were also made of the loss of hydraulic conductivity of twigs subjected to pressure differentials across conduit pit membranes, equalling the leaf water potential at the turgor loss point. The loss of theoretical hydraulic conductivity was calculated as the ratio of i esivr⁴ (where r is the conduit radius) of the non-conducting conduits to that of all the conduits in the outermost wood ring of I, N and J. Stem zones such as 1-year-old nodes and junctions were localized with narrower and shorter xylem conduits and with higher percentages of conduit ends than internodes. Such ‘constricted zonesrsquo; were less vulnerable to embolism than internodes. Latewood conduits were consistently narrower, shorter and less vulnerable to embolism than earlywood ones. A positive relation therefore existed between conduit diameter and length and vulnerability to embolism. The overall vulnerability to embolism of Q. cerris plants is discussed in terms of xylem conduit width and length and of the distribution of conduit ends.     

Hydraulic conductance and rootstock effects in grafted vines of kiwifruit

Whole-plant hydraulic conductance, shoot growth, and leaf photosynthetic properties were measured on kiwifruit vines with four clonal rootstocks to examine the relationship between plant hydraulic conductance and leaf stomatal conductance (gs) and to test the hypothesis that reduced hydraulic conductance can provide an explanation for reductions in plant vigour caused by rootstocks. The rootstocks were selected from four species of Actinidia and grafted with Actinidia chinensis var. chinensis 'Hort16A' (yellow kiwifruit) as the scion. Total leaf area of the scion on the least vigorous Actinidia rootstock, A. kolomikta, was 25% of the most vigorous, A. hemsleyana. Based on shoot growth and leaf area, the selections of A. kolomikta and A. polygama are low-vigour rootstocks, and A. macrosperma and A. hemsleyana are high-vigour rootstocks for A. chinensis. Whole-plant hydraulic conductance, the ratio of xylem sap flux to xylem water potential, was lower in the low-vigour rootstocks, reflecting their smaller size. However, leaf-area-specific conductance (Kl) and gs were both higher in the low-vigour rootstocks, the opposite of the expected pattern. Differences in Kl were found in the compartment from the roots to the scion stem, with no difference between rootstocks in the conductance of stems or leaves of the scion. There was no evidence that the graft union caused a significant reduction in hydraulic conductance of vines with low-vigour rootstocks. Leaf photosynthetic capacity did not vary between rootstocks, but photosynthesis and carbon isotope discrimination (Delta13C) under ambient conditions were higher in the low-vigour rootstocks because gs was higher. gs and Delta13C were positively correlated with Kl, although the mechanism for this relationship was not based on stomatal regulation of a similar xylem water potential because water potential varied between rootstocks. For Actinidia rootstocks, changes in Kl do not provide a direct explanation for changes in vigour of the scion. However, depending on the rootstock in question, changes in hydraulic conductance, biomass partitioning, and crown structure are involved in the response.