The role of carbohydrate reserves in evergreen tree development,with particular reference to macadamia

We take three approaches to more clearly define the role of carbohydrate (CHO) reserves in the development of evergreen trees. First, we examine the lychee and macadamia literature to develop a whole tree carbon budget to show that current photosynthate makes a greater contribution to the carbon for new growth than CHO reserves. Second, we show that the presence of leaves is sufficient for the production of a functional new shoots on small, girdled branches with few CHO reserves. Third, we use a shade experiment to show that short-term suppression of whole-tree photosynthesis can severely affect new shoot development. Subsequent decapitation of all branches on both the shaded and control trees resulted in faster bud release in the shaded trees, notwithstanding lower CHO reserves.     

盐胁迫对果树光合生理的影响

结合我们相关的研究结果综述了盐胁迫对果树光合作用的影响。总体来说盐胁迫导致果树净光合速率下降,且这种作用受树种、品种、砧木以及盐的种类、浓度和处理时期的影响;盐胁迫还使叶片干物质积累下降并影响碳水化合物的组成。盐逆境所产生的渗透胁迫和离子毒害而引起的气孔性限制和非气孔性限制则是果树叶片净光合速率下降的原因。此外,盐胁迫使果树叶片的叶绿素荧光效率下降。文章最后还提了出今后研究果树盐胁迫与光合作用的关系应与实际的生产环境相结合。     

Drying half of the root-zone from mid fruit growth to maturity in ‘Hass’ avocado (Persea americana Mill.) trees for one season reduced fruit production in two years

We tested the effect of extended drying of half the root system on fruit yield and fruit Ca concentration, an indirect measure of fruit quality, in avocado (Persea americana Mill. cv Hass). In a field experiment on a sandy soil, withholding irrigation and plastic sheeting was used to dry the root-zone beneath the whole canopy (DD) or half the canopy (WD), compared with well-watered trees (WW). The irrigation water contained added nutrients and was slightly saline. Yield, shoot growth, leaf conductance, leaf and fruit water status and mineral concentrations of leaves and fruit were studied. The responses of treated trees were assessed in the following season during which normal irrigation practices were restored. With respect to yield, the WD treatment behaved the same as the DD treatment. It reduced yield by more than half and proportionately more than the reduction in water supply thus reducing irrigation efficiency. Re-watering did not restore yield of WD or DD-trees in the next season. The WD and DD treatments had no effect on the concentration of Ca in the fruit mesocarp and so are unlikely to affect fruit quality. The main impact of reduced water supply on the trees was fruit abscission and this was linked to dry soil around the roots rather than the water status of the leaves or fruits. We conclude that extended drying of half of the root-zone in one season reduced irrigation efficiency for two seasons by promoting the abscission of developing fruit to the same extent as occurred when the whole root system was exposed to extended drying.     

果树水分胁迫研究进展

从水分胁迫对果树叶、根的形态指标及显微结构 ,叶片气孔行为、光合作用、光抑制、活性氧代谢、脂氧合酶代谢、多胺代谢、脯氨酸、核酸代谢、内源激素变化等生理生化方面综述了近十几年来的研究成果 ,为全面研究果树抗旱机理及进一步制定抗旱措施奠定理论基础     

Effect of fruit-load on the growth,absorption, and partitioning of inorganic nutrients in young ‘Fuyu’ persimmon trees

Tree growth and the increase in inorganic nutrients during a growing season were studied in 3-year-old, container-grown ‘Fuyu’ persimmon trees with different levels of fruit-load. Fruit-load was varied by adjusting the leaf–fruit ratio to 10, 20, 30, and 60 on July 5. Some trees were completely defruited. The whole trees were harvested on November 5 of that year, and increases in dry weight and inorganic elements in different tree parts during the season were measured. A lower leaf–fruit ratio resulted in a higher yield but also decreased fruit weight, soluble solids, and fruit color. Dry weight of the aerial woods and root increased significantly with a greater ratio. However, total dry weight including the fruits was higher in the lower ratio trees than in the higher ratio trees; the weight of a tree that maintained the ratio of 10 was 35% higher than a tree without fruit. Fruits accounted for 62% of total dry weight when the ratio was 10 and 27% when the ratio was 60. The leaf–fruit ratio did not significantly alter the total increase in P, K, Ca, and Mg content. N tended to decrease with an increasing leaf–fruit ratio. Of the total increases of N, P, and K during the season, the fruits accounted for 30%, 50%, and 54% at the ratio of 20, respectively, and the percentages increased to 44, 81, and 74 at the ratio of 10. The inorganic element content in the aerial woods and root tended to increase with greater ratios, especially in the root. When trees were defruited, N and K increases in the root accounted for 44% and 35% of the total increase, respectively. These results indicated that persimmon orchard fertilization programs with different fruit-load levels should accommodate the proportion of nutrients partitioned to fruits that are removed permanently from the tree.     

Dry weight and carbohydrate distribution in different tree parts as affected by various fruit-loads of young persimmon and their effect on new growth in the next season

This study was conducted to determine the changes in the accumulation of dry weight (DW) and the distribution of carbohydrates in different parts of young persimmon (Diospyros kaki) as affected by various fruit-loads. The effects of such changes were monitored with regard to the abundance of new growth in the following year. On June 15, the fruit-load was adjusted to a leaf–fruit (L/F) ratio of 10, 20, and 30, and some trees were completely defruited. Between June 15 and November 11, the increase in DW was less in the defruited and more in the higher L/F-ratio trees. Among the various tree parts, the DW increased the most in the fruits during the same period. It was observed that the lower the L/F ratio, the more the DW in the fruits: the fruits accounted for 78.7% of the total DW in the 10-L/F, 57.6% in the 20-L/F, and 49.7% in the 30-L/F ratio trees. In contrast, as the L/F ratio increased, the DW distribution to the roots increased to 3% in the 10-L/F, 13% in the 20-L/F, and 26% in the 30-L/F ratio trees and 61% in the defruited trees. During this period, carbohydrates were distributed mostly to the fruits, but as the L/F ratio decreased, their distribution to permanent tree parts decreased. The distribution of soluble sugars and starch to the permanent parts was 1% and 14% in the 10-L/F, 3% and 48% in the 20-L/F, 9% and 57% in the 30-L/F ratio trees, and 71% and 93% in the defruited trees, respectively. Maintaining a lower L/F ratio resulted in a decrease in the shoot and fruit numbers in the following year: there were zero and four fruits in those trees with an L/F ratio of 10 and 20, respectively. The carbohydrates decreased in the roots of both the 30-L/F ratio trees and the defruited trees, whereas the defruited trees showed more new shoot growth in the following year. In the roots of the trees with the L/F ratio of 30, soluble sugars and starch decreased by 7.6 g and 1.8 g, respectively, during the spring growth, whereas, in the roots of the defruited trees soluble sugars and starch decreased by 33.3 g and 94.6 g, respectively.     

螺旋环剥对‘糯米糍’荔枝坐果与碳水化合物及激素的影响

研究结果表明，幼‘糯米滋’荔枝主要螺旋环剥能减少碳水化合物向根系转运，从而增加叶片和果实中的碳水化合物含量，增多果实中赤霉素类物质和降低脱落酸类物质含量。坐果率的提高与赤霉素／脱落酸的比值较高关系密切。     

Studies on the relationship between leaf number and area and fruit development in mango (Mangifera indica L.)

The optimum leaf number: fruit ratio in various mango cuitivars was sought by isolating individual fruits with known numbers of supporting leaves by shoot girdling. ¹⁴CO₂ feeding experiments showed a higher rate of carbon fixation in the leaves of girdled shoots than of control shoots but the translocation of ¹⁴C assimilates to the developing fruits on the girdled and control shoots was comparable. Starch accumulation in the leaves was reduced by shoot girdling. The stomatal resistance of the leaves of girdled shoots was comparable to that of leaves on control shoots. In all the cuitivars studied it was observed that 30 leaves, the maximum available on a shoot, could not support the growth of a single fruit to normal size. The results also show that fruit development depends not only on the current assimilates but also to a great extent on reserves. The utilization of reserve metabolites from vegetative organs during the ‘on’ year could be a contributing factor towards biennial and erratic bearing.     

Status of nitrogenous and carbohydrate compounds as affected by nitrogen fertigation rates in young persimmon trees

The effects of nitrogen (N) fertigation rates on levels of nitrogenous and carbohydrate compounds were determined in young ‘Fuyu’ persimmon trees. Total N amounts of 0 g, 10 g, 20 g, 30 g, and 40 g were distributed, respectively, to a 50-L pot by fertigation as different volumes of 1% (w/v) urea solution every week from May 15 to September 25. Fruits were sampled on November 2, and shoots and roots on November 17. High N fertigation rates significantly increased the N and amino acids in all the tree parts and protein in fruits and shoots. In contrast, the concentration of soluble sugars significantly decreased in the fruits and perennial parts as the N rate increased from 10 g to 40 g. A notable reduction of starch was also found in the shoots and roots of trees with high N fertigation rates. The percentage of shoots died due to cold injury during the winter was the highest at the 0 g N rate; however, it was 2.3-fold greater at the 40 g N rate as compared to the 10 g N rate. We concluded that high N fertigation could interfere not only with sugar accumulation of fruits through increased N metabolism, but also reserve carbohydrates of perennial organs.     

Response of evergreen perennial tree crops to gibberellic acid is crop load-dependent. I: GA3 increases the yield of commercially valuable ‘Nules’ Clementine Mandarin fruit only in the off-crop year of an alternate bearing orchard

Worldwide, gibberellic acid (GA₃) is used routinely to increase fruit number and size of seedless mandarins. The efficacy of seven combinations of GA₃ concentrations and application times to maximize total yield and yield of commercially valuable fruit (diameter 57.2–76.2 mm) of ‘Nules’ Clementine mandarin (Citrus reticulata Blanco) was determined in a commercial orchard. GA₃ applied during the period of intense flower abscission failed to reduce the total number of abscised flowers in both the light off- and heavy on-bloom years. No GA₃ treatment reduced fruit abscission when trees were setting the low yield off-crop. However, all trees receiving GA₃ in the high yield on-crop year had fewer abscised fruit than untreated control trees (P = 0.0188) and GA₃ applied 10 days after 75% petal fall and in July increased the number of fruit retained on tagged branches >20% compared to control trees (P = 0.0005). Maximum air temperature was not related to flower or fruit abscission. In the off-crop year (548 fruit per untreated control tree), it was necessary to apply 15 or 25 mg L⁻¹ GA₃ at 60% bloom, 90% bloom, 75% petal fall and 10 days after 75% petal fall to significantly increase the number of fruit per tree and yield of commercially valuable fruit (kilograms and number per tree) (P < 0.0001) above that of control trees, with no reduction in total kilograms per tree. In the following on-crop year, it was better not to apply GA₃: no treatment increased total yield or fruit size and five of seven GA₃ treatments tested reduced total yield as kilograms and number of fruit per tree (P = 0.0003). The results provide strong evidence that GA₃ efficacy is crop load-dependent and dictate that crop load should be considered when using GA₃ to increase fruit set or fruit size of mandarins.     

螺旋环剥对幼龄‘桂味’荔枝果期光合和蒸腾作用的影响

以6年生幼龄‘桂味’荔枝(Litchi chinensis Sonn.‘Guiwei’)为试材,于5月果实发育期,从螺旋环剥处理和对照树上分别选取来自同一基枝的有果枝和无果枝,观察枝梢生长的情况并进行叶片光合和蒸腾作用的研究。结果表明:螺旋环剥有利坐果;螺旋环剥缩小有果叶与无果叶在光合效率上的差别,显著降低了叶片的最大光合速率(Amax)、表观量子效率(AQY)和羧化效率(CE),提高了光补偿点(LCP),削弱了光合效率;净光合速率(Pn)与蒸腾速率(Tr)极显著相关,螺旋环剥显著降低了叶片的Tr,Pn也下降但差异不显著,蒸腾作用减弱的程度比光合作用减弱的程度大;对照叶片的Pn和Tr日变化为单峰曲线型,14:00时达最高峰,螺旋环剥叶片的Pn和Tr在13:00时有明显的午休现象,气孔限制是午休的主要因素;相同处理的有果枝叶片,其日均Pn和Tr都高于无果枝叶片,说明果实的存在可提高‘桂味’荔枝的Pn和Tr。     

Investigations on the use of (2-chloroethyl) phosphonic acid (ethephon,CEPA) for the control of biennial bearing in mango

Biennial bearing is a serious economic problem facing mango growers. In the past it has been attributed to the lack of new growth and insufficient carbohydrate and nitrogenous reserves following a year of heavy cropping. The results of our studies conducted over a period of 4 years have shown that ethephon (200 ppm) could be used to induce heavy flowering and fruiting during the “off year” in the notedly biennial bearing cv. ‘Langra’. The spray applications of ethephon should be started during September and continued four to five times at 15–20 days interval. Higher concentrations of ethephon, viz. 500, 1000 and 2000 ppm, induced moderate to heavy leaf abscission in the experimental trees. Consecutive applications of 200 ppm ethephon for a period of 3 years did not show any decline in the vigour and yielding ability of the treated ‘Langra’ mango trees.     

Timing of fruit removal affects concurrent vegetative growth and subsequent return bloom and yield in olive (Olea europaea L.)

Olive (Olea europaea) demonstrates a high tendency toward alternate fruit production, with significant negative consequences on the industry. Fruit load is one of the main cause-and-effect factors in the phenomenon of biennial bearing, often disrupting the balance between reproductive and vegetative processes. The objectives of the present study were to identify the time range during which heavy fruit load reversibly interrupts the reproductive processes of the following year. The linkage between timing of fruit removal, vegetative growth, return bloom, and fruit yield was studied. Complete fruit removal in cv. Coratina until about 120 days after full bloom (August 15) caused an immediate resumption of vegetative growth. The new shoots grew to twice the length of those on trees that underwent later fruit removal. Moreover, a full return bloom, corresponding with high subsequent yields, was obtained by early fruit removal, while poor or no bloom developed on late-defruited or control trees. Thus, the critical time to affect flowering and subsequent fruiting in the following year by fruit thinning occurs in olive trees even weeks after pit hardening—much later than previously suggested. Furthermore, the data indicate that flowering-site limitation, due to insufficient or immature vegetative growth during the On-year, is a primary factor inducing alternate bearing in olive.     

Mitigation of severe water stress by fruit thinning in ‘O’Henry’ peach: Implications for fruit quality

Fruit thinning can be used to relieve water stress in peach but it is not known how fruit quality will respond under water limited conditions. To elucidate this, we applied, over the growing seasons of 2003 and 2004, irrigation and fruit thinning treatments to ‘O’Henry’ peach at the onset of Stage III of fruit development. The treatments were full irrigation (FI) and no irrigation (NI) with each receiving three crop load levels: light, commercial and heavy. Fruit size and fruit skin colouration were lower in NI than in FI but the following were higher in NI fruit: firmness, juice titratable acidity and electrical conductivity, and concentrations of dry matter and soluble solids. All the quality parameters were significantly correlated with tree water status, expressed as midday stem water potential. NI fruit matured 5–10 days later than FI fruit. The increased dry matter concentration, juice acidity, and electrical conductivity in NI fruit were indicative of lower quality at harvest. Choosing a light crop load was not effective in improving NI fruit quality except for the size. Since fruit size is the most important fruit quality attribute, its increase might compensate for yield losses resulting from heavy fruit thinning. Nevertheless, when heavy thinning is performed for mitigation of water stress in Stage III, the overall fruit quality will suffer if severe water stress is inevitable because of shortage or lack of irrigation water.     

茭白碳水化合物积累与分配特性研究

 以茭白品种‘蒋墅茭’和‘葑红早’为试材的研究表明: 在肉质茎膨大之前, 植株茎蘖各部位的总糖、还原糖、淀粉积累量持续上升, 短缩茎中积累量最高; 肉质茎膨大开始时, 各部位碳水化合物积累量下降, 以短缩茎的下降量最大, 认为膨大前叶片的光合产物在短缩茎和叶片中积累以短缩茎为主,植株积累的碳水化合物主要以淀粉形式存在。孕茭前¹⁴C 同化物在短缩茎的分配率较大, 上位叶片、叶鞘也有较多积累, 肉质茎膨大过程中, 大量的¹⁴C 同化物在肉质茎中积累。饲喂叶片光合产物除供应自身茎蘖外, 还向植株其它各部位输送。     

Effects of shading and thidiazuron + oil treatment on dormancy breaking,blooming and fruit set in apricot in a warm-winter climate

Overcoming dormancy represents one of the major limitations to fruit tree production in warm areas. High temperatures during the chilling period have a negative effect on breaking of dormancy. Shading of trees reduces the incidence of radiation and the temperature. Previous works on shading did not take into account the stage of dormancy. The aim of this work was to evaluate the effects of shading during different periods of endodormancy, in an area having relatively warm winters, and the effects of a treatment of thidiazuron ([TDZ] N-phenyl-N-1,2,3-thiodiazol-5-il-urea) and winter oil, on apricot blooming, fruit set and ripening. The study was carried out during three years which showed marked differences regarding chilling accumulation. Autumn shading did not affect blooming or harvesting. Regarding harvest date, 2–3 days of precocity were achieved by shading during endodormancy, relative to the control. The TDZ + oil treatment increased blooming percentage, made blooming more uniform and brought forward the blooming (by 7–14 days) and ripening (3–8 days) dates. However, pistil abortion percentage was strongly increased by using TDZ and winter oil when there was low chilling accumulation, which led to reduce fruit set percentage. Shading during late endodormancy produced 5 days precocity for the harvest date in the year with lower chill accumulation. Shading of trees during endodormancy and TDZ + oil treatment could be suitable for increasing precocity in warm-winter climates. Significant year-to-year variation has been shown for blooming, pistil abortion, fruit set and fruit/bud percentages. Correlations among these variables are also discussed. Under conditions of marginal chill accumulation, the distribution of chilling during autumn and winter had an important role in the overcoming of dormancy, blooming and fruit set.     

Content levels of various fruit metabolites in the ‘Conference’ pear response to branch bending

The response of ‘Conference’ pear subjected to branch bending in the content levels of various metabolites in its fruit was investigated. The fruits in commercial maturity were sampled in 2004 and 2005 from branches bent in the late summer of 2003 (the summer treatment), from branches bent in the late spring of 2004 (the spring treatment) and from control branches. The content levels of carbohydrates, organic acids and phenolic compounds were compared among treatments in two successive years. The fruit revealed various responses in content levels of metabolites. In the first year after bending, no significant differences were found in the content of each carbohydrate, but in 2005 by far the highest content level of glucose and fructose and the lowest content level of sucrose were found in fruits from the control. The control fruit showed significantly higher content levels of malic acid and lower content levels of some phenolics (chlorogenic and vanillic acid) in the first year after bending, but in the next year the opposite reaction occurred—the control fruit had the lowest content level of malic acid and the highest content level of epicatechin, quercetin-3-d-galactoside and quercetin-3-β-d-glucoside. The comparison of the two bending treatments alone in 2004 showed that the summer treatment often produced a slightly higher value of each phenolic in comparison to the spring treatment. However, in 2005 the significantly highest content of chlorogenic acid was in fruit from the spring treatment. Sorbitol, as well as citric acid, catechin and sinapic acid showed no clear tendency among treatments, neither in 2004 nor in 2005. It is suggested that these variations of ‘Conference’ fruit subjected to different bending treatments could not be the result of bending alone, but that they could be indirectly affected by other physiological responses of the fruit tree. However, it seemed that variations are affected by the time of bending and by the year-to-year, and such responses can be attributed to the ‘Conference’ genotype only.     

不同物候期枣树细根和叶片可溶性糖、蛋白和C、N、P含量的变化

以陕北‘七月鲜’枣树为材料,研究不同物候期(萌芽期、展叶初期、展叶中期、全叶期、果实成熟期、落叶期)细根和叶中可溶性糖、蛋白以及C、N、P的含量变化规律。结果表明:(1)随着根序增大,细根中可溶性糖、蛋白、有机C含量均呈增加趋势,全N和全P含量呈降低趋势;(2)不同物候期细根中可溶性糖和蛋白含量呈"W"变化趋势,叶中呈先上升后下降的态势;(3)细根的有机C含量呈先降低后增加的趋势,最低值出现在果实成熟期,叶呈先降低后增加再降低的趋势,最低值出现在展叶中期;(4)细根的全N含量呈先降低后增加的趋势,叶呈先降低后增加再降低,在全叶期和落叶期含量较低;(5)细根的全P含量呈先降低后增加再降低的趋势,叶持续降低,直到落叶期有所增加;(6)细根与叶片的可溶性糖、可溶性蛋白和C、N、P之间的相关性不显著,细根与叶片的各指标之间相互关系有待进一步研究。     

Root influence on shoot development in tea (Camellia sinensis (L.) O. Kuntze) following shoot pruning

The physiological interaction between the factors controlling the development of new shoots in pruned tea plants was investigated in an experiment using a widely cultivated TRI clone growing at low elevation in Sri Lanka. The functional importance during new shoot growth of root reserves, feeder roots and leaves and buds on unpruned branches (lung-shoots) is critically evaluated. A scheme for their interaction is presented, which highlights the need to maintain feeder roots alive during new shoot development, for which a large proportion of both reserve and current assimilates seem to be utilized. Mature leaves on lung-shoots, besides supplementing carbohydrate supply by current photosynthesis, may also serve to maintain the flow of metabolites from feeder roots to developing shoots and also function as ‘sinks’ for excess, toxic levels of root metabolites. Removing all mature leaves, as is done during ‘clean pruning’, leads to the depletion of reserves below the critical level for feeder root survival, resulting in death of feeder roots and failure of new buds to form shoots.     

去果对桃源叶碳水化合物含量及其相关酶活性日变化的影响

 研究了‘艳丰1号’桃果核硬化期和果实最后迅速生长期去果对源叶碳水化合物含量及其相关酶活性日变化的影响。去果处理和留果对照源叶中总可溶性糖、山梨醇、蔗糖和淀粉含量均持续升高,而葡萄糖和果糖含量持续降低。在全天大多数时间里去果处理源叶中山梨醇和淀粉含量及它们的关键合成酶6 - 磷酸醛糖还原酶(A6PR) 、腺苷二磷酸葡萄糖焦磷酸化酶(ADPGPPase) 活性均显著高于留果对照。就库源关系中可溶性糖和淀粉积累对光合作用的调控进行了讨论。