Effect of fruiting on carbon budgets of apple tree canopies

Summary Carbon budgets were calculated from net photosynthesis and dark respiration measurements for canopies of field-grown, 3-year-old apple trees (Malus domestica Borkh.) with maximum leaf areas of 5.4 m² in a temperature-controlled Perspex tree chamber, measured in situ over 2 years (July 1988 to October 1990) by computerized infrared gas analysis using a dedicated interface and software. Net photosynthesis (Pn) and carbon assimilation per leaf area peaked at respectively 8.3 and 7.7 mol CO₂ m^–2 s^–1 in April. Net photosynthesis (Pn) and dark respiration (Rd) per tree peaked at 3.6 g CO₂ tree^–1 h^–1 (Pn) and 1.2 g CO₂ tree^–1 h^–1 (Rd), equivalent to 4.2 mol CO₂ (Pn) and 1.4 mol CO₂ (Rd) m^–2 s^–1 with maximum carbon gain per tree in August and maximum dark respiration per tree in October 1988 and 1989. In May 1990, a tree was deblossomed. Pn (per tree) of the fruiting apple tree canopy exceeded that of the non-fruiting tree by 2–2.5 fold from June to August 1990, attributed to reduced photorespiration (RI), and resulting in a 2-fold carbon gain of the fruiting over the non-fruiting tree. Dark respiration of the fruiting tree canopy progressively exceeded, with increasing sink strength of the fruit, by 51% (June–August), 1.4-fold (September) and 2-fold (October) that of the non-fruiting tree due to leaf (i. e. not fruit) respiration to provide energy (a) to produce and maintain the fruit on the tree and (b) thereafter to facilitate the later carbohydrate translocation into the woody perennial parts of the tree. The fruiting tree reached its optium carbon budget 2–4 weeks earlier (August) then the non-fruiting tree (September 1990). In the winter, the trunk respired 2–100 g CO₂ month^–1 tree^–1. These data represent the first long-term examination of the effect of fruiting without fruit removal which shows increased dark respiration and with the increase progressing as the fruit developed.     

Effects of defruiting on source-sink relationship, carbon budget, leaf carbohydrate content and water use efficiency of apple trees

Three-year-old apple ( Malus domestica Borkh. cv. Golden Delicious) trees with ca 5 m² leaf area each were grown in situ in lysimeters and defruited either in August. September or October. Net photosynthesis, dark respiration and water use efficiency were measured for 5 days prior to and after fruit removal. using tree canopy chambers and infrared gas analysis with a dedicated interface and software. Net photosynthesis of the tree canopy declined after defruiting. irrespective of the month of defruiting. The decline after fruit removal was 30-40% in August or September and 57% in October. After defruiting in August or September, trees respired 50–60% slower in the dark, but 39% faster after defruiting in October. The excessive respiratory losses after fruit removal in October. when the tree lost more carbon than it assimilated. may have been induced by translocation of carbohydrates from the leaves to the perennial woody parts of the tree and by the onset of leaf senescence. Fruit removal in August or September increased water use efficiency by 31–41%. but decreased water use efficiency in October by 75%     

Small fruit problem in Citrus trees

Summary Physiological causes of the small fruit problem which occurs in certain trees of orange [Citrus sinensis (L.) Osbeck cv. Valencia] were investigated in terms of water relations and gas exchange of fruits during early fruit development as well as tree carbohydrate reserves. These data from cv. Valencia trees with and without a small fruit potential were compared with those of the large fruited cv. Navel. Neither fruit water potential nor fruit transpiration nor tree carbohydrate reserves appeared to be a cause of the small fruit. Yield records showed the small fruit to be assocaited with a large number of fruit per tree. However, fruits from cv. Valencia trees with a small fruit potential respired faster than either fruits of the same cultivar and size from trees without the physiological disorder or fruits of the same size of cv. Navel and also exceeded the dark respiration of the respective leaves. Hence, the small fruit problem in cv. Valencia was partly attributed to inefficient fruit photosynthesis, causing excessive respiration of each of a larger number of fruits compared to fruits of a tree of the same cultivar but without the physiological disorder. Fruits of cv. Valencia respired more in their 2 months longer lifetime on the tree relative to those of cv. Navel. It is concluded that orchard management methods will have to be investigated to balance the fruit load of the cv. Valencia tree utilizing the carbon available for fruit growth and to minimise stress during the early fruit development.     

间伐改形对陇东旱塬密闭苹果园树体冠层结构和发育后期叶片质量的影响

为研究间伐改形对红富士成龄乔化密植果园树体冠层特征、生育后期叶片生理特性、养分积累分配规律、光合生产力和土壤水分时空分布动态的影响,以18年生‘红富士’苹果密闭园为试材,对苹果树单株结构参数、枝量、枝类组成、覆盖率、叶片光合速率等参数进行测定.结果表明: 密闭果园叶片光合作用受非气孔因素限制,导致PSII最大光化学效率、PSII光合潜能、光合性能指数下降了1.2%、11.5%、13.9%.间伐改形后,叶面积指数、树冠覆盖率有所降低,使得冠层直射光透过系数增加了79.0%,树形结构有所改善.苹果园总枝数降低到约1100400条·hm^-2,单株枝量增加了5.0%,短枝比例提高至73.0%.由于冠层光照条件的改善,叶面积、比叶质量、百叶重、叶绿素含量有不同程度的提高.叶片光合速率的提高促进了光合产物的积累,淀粉、蛋白质含量为密闭果园的143.5%、107.8%.叶片的发育质量与其所处的光照辐射环境有着密切联系,密闭果园经间伐、改形后,果园群体结构和冠层光照得以改善,促进了叶片生长发育,提高了叶片光合效能,降低了果园土壤水分的无效消耗,是陇东旱塬苹果产区密闭果园适宜的调整、优化方案.     

Seasonal Changes in the Photosynthetic Rate in Apple Trees : A Comparison between Fruiting and Nonfruiting Trees

Seasonal changes in photosynthesis of apple trees (Malus domestica Borkh.) were monitored to examine the effect of source-sink interactions on photosynthesis and photorespiration. Elevated photosynthetic rates were observed during two periods of the growing season and correlated with the fruiting process. The first period of increased photosynthetic rates was during the bloom period, when spur leaves on flowering shoots exhibited up to 25% higher photosynthetic rates than vegetative spur leaves on a leaf area basis. CO₂ assimilation rates were also higher in fruiting trees than nonfruiting trees during the period of rapid fruit growth from July to September. Photorespiration, dark respiration, leaf resistance, and transpiration exhibited no seasonal changes which correlated to the presence or absence of fruit. These data represent the first comprehensive examination of the effects of flowering/fruit formation on photosynthesis and photorespiration in perennial plants.     

Influence of Seasonal Variations in Soil Water Availability on Gas Exchange of Tropical Canopy Trees

Seasonal variations in environmental conditions influence the functioning of the whole ecosystem of tropical rain forests, but as yet little is known about how such variations directly influence the leaf gas exchange and transpiration of individual canopy tree species. We examined the influence of seasonal variations in relative extractable water in the upper soil layers on predawn leaf water potential, saturated net photosynthesis, leaf dark respiration, stomatal conductance, and tree transpiration of 13 tropical rain forest canopy trees (eight species) over 2 yr in French Guiana. The canopies were accessed by climbing ropes attached to the trees and to a tower. Our results indicate that a small proportion of the studied trees were unaffected by soil water depletion during seasonal dry periods, probably thanks to efficient deep root systems. The trees showing decreased tree water status (i.e., predawn leaf water potential) displayed a wide range of leaf gas exchange responses. Some trees strongly regulated photosynthesis and transpiration when relative extractable water decreased drastically. In contrast, other trees showed little variation, thus indicating good adaptation to soil drought conditions. These results have important applications to modeling approaches: indeed, precise evaluation and grouping of these response patterns are required before any tree‐based functional models can efficiently describe the response of tropical rain forest ecosystems to future changes in environmental conditions.     

Photosynthetic responses of Miconia species to canopy openings in a lowland tropical rainforest

We examined the photosynthetic acclimation of three tropical species of Miconia to canopy openings in a Costa Rican rainforest. The response of photosynthesis to canopy opening was very similar in Miconia affinis, M. gracilis, and M. nervosa, despite differences in growth form (trees and shrubs) and local distributions of plants (understory and gap). Four months after the canopy was opened by a treefall, photosynthetic capacity in all three species had approximately doubled from closed canopy levels. There were no obvious signs of high light damage after treefall but acclimation to the gap environment was not immediate. Two weeks after treefall, A_max, stomatal conductance, apprarent quantum efficiency, and dark respiration rates had not changed significantly from understory values. The production of new leaves appears to be an important component of light acclimation in these species. The only variables to differ significantly among species were stomatal conductance at A_max and the light level at which assimilation was saturated. M. affinis had a higher stomatal conductance which may reduce its water use efficiency in gap environments. Photosynthesis in the more shade-tolerant M. gracilis saturated at lower light levels than in the other two species. Individual plant light environments were assessed after treefall with canopy photography but they explained only a small fraction of plant variation in most measures of photosynthesis and growth. In conclusion, we speculate that species differences in local distribution and in light requirements for reproduction may be more strongly related to species differences in carbon allocation than in carbon assimilation.     

Carbon balance in a monospecific stand of an annual herb <Emphasis Type="Italic">Chenopodium album</Emphasis> at an elevated CO<Subscript>2</Subscript> concentration

Elevated CO₂ enhances carbon uptake of a plant stand, but the magnitude of the increase varies among growth stages. We studied the relative contribution of structural and physiological factors to the CO₂ effect on the carbon balance during stand development. Stands of an annual herb Chenopodium album were established in open-top chambers at ambient and elevated CO₂ concentrations (370 and 700 μmol mol⁻¹). Plant biomass growth, canopy structural traits (leaf area, leaf nitrogen distribution, and light gradient in the canopy), and physiological characteristics (leaf photosynthesis and respiration of organs) were studied through the growing season. CO₂ exchange of the stand was estimated with a canopy photosynthesis model. Rates of light-saturated photosynthesis and dark respiration of leaves as related with nitrogen content per unit leaf area and time-dependent reduction in specific respiration rates of stems and roots were incorporated into the model. Daily canopy carbon balance, calculated as an integration of leaf photosynthesis minus stem and root respiration, well explained biomass growth determined by harvests (r ² = 0.98). The increase of canopy photosynthesis with elevated CO₂ was 80% at an early stage and decreased to 55% at flowering. Sensitivity analyses suggested that an alteration in leaf photosynthetic traits enhanced canopy photosynthesis by 40–60% throughout the experiment period, whereas altered canopy structure contributed to the increase at the early stage only. Thus, both physiological and structural factors are involved in the increase of carbon balance and growth rate of C. album stands at elevated CO₂. However, their contributions were not constant, but changed with stand development.     

Community homogenization and the invasiveness of commensal species in Mediterranean afforested landscapes

The ecological consequences of homogenization remain relatively unexplored. One example of landscape-homogenizing is the establishment of plantations. We studied the effect of human-made forests by contrasting plant and small-mammal community composition between planted tree stands and adjacent natural habitat in two different Mediterranean habitats in Israel: (1) inland habitat where we focused on pine (Pinus halepensis) and carob (Ceratonia siliqua) stands, and (2) coastal sand dune habitat where we focused on planted acacia (Acacia saligna) stands. We first wanted to verify whether planted trees modify plant species composition, and second, if and how the small-mammal community is affected by the different habitat conditions created in plantations with different canopy cover. We were especially interested in the abundance of the commensal house mouse (Mus musculus). All tree stands underwent biotic homogenization indicated by abundance of house mice coupled with lower diversity of indigenous vegetation and small-mammal abundances and diversities. Habitat structural diversity was positively related with small-mammals diversity and was lower in artificial tree stands in both habitats. Our results suggest that using the abundance of commensal generalist species such as the house mouse relative to other more specialist small-mammals is a good approach to determine ecosystem integrity. Pre-commercial thinning treatment is a potential management tool to maintain a proportion of native tree species within the canopy of planted tree stands. However, until sufficient data is available for making generalizations, the exact level of thinning necessary to reverse the homogenization processes in man-made plantations and keeping indigenous small-mammal communities diverse and less prone to invasion must be determined empirically.     

Evaluation of monoammonium phosphate and bacterial strains to increase tree growth and fruit yield in apple replant problem soil

The long-term effects of biological agents alone and in combination with monoammonium phosphate on tree growth and fruit production of apple trees planted on apple replant soil was studied for five years. Application of monoammonium phosphate (MAP) in the year of planting increased shoot growth, cross-sectional trunk area and fruit yield of McIntosh on M.26 rootstock for the first two years. The application of bacterial agents alone were not effective in increasing young tree growth except BACT-1 in 1987. None of the bacterial agents increased fruit yield when applied alone. The addition of certain bacterial agents to MAP application increased young tree growth in various years. The combination of bacterial agent B-10 and MAP reduced young tree growth and yield compared with the MAP treatment alone. These results suggest that the application of MAP alone may be sufficient to alleviate the replant problem and the addition of BACT-1, EBW-4 or B8 bacterial agents to this treatment may be beneficial to increase tree growth in some years. Contribution number 822. Contribution number 822.     

矮嵩草草甸植物群落的光合特性研究

 矮嵩草草甸植物群落的光合、暗呼吸和土壤呼吸的研究表明：光合作用的日变化在6月份接近平坦型,7、8月份呈午间降低型。矮嵩草草甸植物群落的光合作用受较低的光合面积指数及冠层叶片的受光势态的影响,存在着明显的光饱和现象,光补偿点及光饱和点相对于全日照光合有效辐射均较低,接近于单叶的光响应特性。裸露地面的土壤呼吸和植物与土壤体系的暗呼吸不仅与温度有关,而且与土壤水分状况和降雨量也有密切联系。影响草甸群落光合特性的主要因素有：高原地区强烈的太阳辐射,较低的光合面积指数和植物根系与土壤紧密结合所形成的草结皮层结构。     

疏花对富士苹果光合同化物分配的外部调节和生理表现

在盛花期,利用对树冠不同部位疏除花序的试验。对富士苹果树体内同化物的调配及其生理表现进行了研究。结果表明,苹果平均单果重和平均单果叶面积之间关系呈线性正相关。该相关关系在不同处理之间无差异（r=0.91),充分说明在苹果树体内存在一个可自由流动的碳水化合物库或者说树体对结果具有光合调节作用,疏花可以从外部调节光合同化物分配的方向。对每株苹果树未结果部位和结果部位短枝及延长枝叶片净光合速率测定表明。结果和未结果部位光合速率没有显著差异。虽然疏花没有显著影响苹果单株枝条生长量和叶面积,但是,苹果树树冠全部疏除花序的一侧 或主枝的平均枝条生长量和叶面积去比该树其它结果部位分别平均高出153.5cm和8900cm^2,差异极为显著。苹果树疏除花序1/2或3/4的数量使苹果平均单果重比不疏花分别显著增加了33g或79g。而果实钙浓度每百克鲜重分别降低了1.62mg或2.66mg,充分说明当疏花显著地影响果实大小时,果实内钙浓度也会明显地发生相应的变化,虽然在疏花处理之间不同部位疏除花序没有影响单株花芽总数量,但是,每株树前一年春季疏花的部位比未疏花,结果的部位极显著地增加了花芽数量（SED=11.6)。     

Effect of plant architecture and hail nets on temperature of codling moth habitats in apple orchards

Plant architecture of apple trees in commercial orchards was rapidly changed from traditional tall trees to dwarf trees to optimize yield and fruit quality. Additionally, hail nets are widely used to prevent yield loss by hail. These changes are expected to considerably influence the orchard microclimate and thus the developmental rates of pest insects in apple. However, these relationships have not yet been fully elucidated. The present study was conducted over the seasonal cycle to investigate the influence of plant architecture and hail nets on the habitat temperatures of the codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), in apple, Malus domestica Borkh. (Rosaceae). Within the canopies, leaf area index (LAI) and global site factor (GSF) were quantified using hemispherical photography. Temperature was analysed for the main habitats of the different codling moth stages, i.e., air within the canopy, bark of tree stems, and apple fruit. In dwarf trees, LAI was lower, leading to a higher GSF than in tall trees. Hail nets did not influence LAI and GSF. Results for dwarf trees compare as follows with those for tall trees: Average air temperatures within the canopy were 0.7 °C higher during daytime, whereas 0.4 °C lower at night. Mean surface temperatures of bark were 0.9 °C higher on sunny and 0.4 °C on overcast days. Mean surface temperatures of apple fruits were 1.8–2.7 °C higher on sunny days, but 0.6 °C cooler on overcast days. The effect of hail nets was confined to a reduction of the air temperature within the canopy by approximately 0.2–0.8 °C. Bark and apple surface temperatures were not significantly affected. Based on the temperature differences in the habitats considered, the calculated development of the codling moth in dwarf trees was on average 3 days faster than in tall trees. The calculations imply a negligible effect of hail nets on codling moth development.     

Host marking pheromone of Rhagoletis cerasi: field deployment of synthetic pheromone as a novel cherry fruit fly management strategy

We tested the efficacy of synthetic host marking pheromone (HMP) of the European cherry fruit fly (R. cerasi) as a fruit-infestation-reducing-agent in an experimental cherry orchard. Two different pheromone deployment strategies were compared: covering entire tree canopies with synthetic HMP or treating only one half (top to bottom or lower half) of the tree canopy. Pheromone application caused a tenfold reduction in fruit infestation if the entire tree canopy was covered (0.226 vs 0.021 pupae/fruit in untreated and treated trees, respectively). Results show, nevertheless, that a significant reduction in fruit infestation can be achieved by treating only one half of tree canopies (top to bottom) (0.021 vs 0.048 pupae/fruit when comparing totally vs partially treated trees). We conclude that synthetic cherry fruit fly HMP has potential as a fruit fly management tool, especially in cases where organically grown fruit reaches high market values.     

Size of orchard trees as a factor affecting behavioural control of apple maggot flies (Dipt., Tephritidae) by traps

Over a 3-year period (1997–1999), we examined the influence of tree size on effectiveness of traps for behaviourally controlling apple maggot flies, Rhagoletis pomonella (Walsh), in small blocks of trees in Massachusetts commercial apple orchards. Traps were red spheres coated with Tangletrap and baited with an attractive synthetic host fruit odour (butyl hexanoate). Traps were placed 6 m apart on perimeter apple trees of each block and were designed to intercept apple maggot flies immigrating into blocks from unmanaged host trees. Based on captures of adults by unbaited red spheres placed near the centre of each block to monitor degree of adult penetration into the interior and on percentages of fruit injured by apple maggot, traps surrounding blocks of small trees (1.5 m canopy diameter) planted at high density were more effective in controlling apple maggot flies (relative to control obtained by insecticide sprays used in comparison blocks) than were traps surrounding blocks of large trees (3.7 m canopy diameter) planted at low density, with traps surrounding blocks of medium-size trees (2.5 m canopy diameter) planted at medium density generally providing an intermediate level of control.     

Evaluation ofMicrothrix inconspicuella, [Lepidoptera: Pyralidae], a potential biological control agent forEmex australis in Australia,carried out in apple orchards in South Africa

Host specificity tests carried out in the laboratory in Australia during 1977, showed thatMicrothrix inconspicuella Ragonot could develop on young apple leaves (Harley et al., 1979). Field studies in unsprayed apple orchards in South Africa showed that some feeding occurred, but fewer than 40% of late instar larvae developed to adults when confined in sleeves on apple tree branches. No feeding or survival occurred in large field cages or in the open. Adults which developed from apple fed larvae were smaller, deformed, occasionally mated and laid fertile eggs but their progeny did not feed or develop on apple fruit or leaves. In conclusion,M. inconspicuella larvae did not develop on apple fruit or leaves in the field, damage was mainly limited to apples already injured and feeding on leaves was minimal. Under normal pest control practicesM. inconspicuella populations did not survive on any part of the apple tree or onE. australis growing under the trees.      

Variation of ginkgolides and bilobalide contents in leaves and cell cultures of<Emphasis Type="Italic">Ginkgo biloba</Emphasis> L.

Ginkgolides (GK) and bilobalide are valuable compounds that belong to the lactone terpene. The contents of these metabolites were determined by HPLC from female and male tree ofGinkgo biloba L. The productivity ofG. biloba cells was also compared with the corresponding individual trees. High variations in the ginkgolides and bilobalide were observed from different individuals, plant parts, and cultured cells. The ginkgolides and bilobalide contents were different depending on the plant parts. Callus was obtained from various plant tissues, and NAA was better at callogenesis than 2,4-D in both the female and male trees. The plants and their corresponding cells showed considerable variation in their ginkgolides and bilobalide concentrations. The ginkgolides and bilobalide contents were not correlated with the production between dominant trees and their corresponding cells. Light irradiation enhanced the production of GK-A and GK-B, however, the concentration of bilobalide decreased under dark conditions.     

Remarks on mixotrophic and autotrophic carbon nutrition of Vitis plantlets cultured in vitro

Two Vitis species were cultured in vitro under photoautrophic (sucrose-free culture medium) and photomixotrophic (sucrose 15 g l^-1) conditions during the period following microcutting rooting (day 34 to day 120). Several parameters were measured at the end of the culture: growth, plant dry weight, carbohydrate uptake from the medium and rates of photosynthesis and dark respiration. The two species behaved very differently. Under photoautotrophic conditions, dark respiration, net photosynthesis and daily CO₂ fixation were higher in Vitis vinifera than in Vitis rupestris. Culture under mixotrophic conditions caused increase in growth, respiration and photosynthesis in Vitis rupestris. In contrast, photosynthesis decreased in Vitis vinifera under the same conditions.     

Developments in the chemical thinning of apple and pear

A literature review is presented on past and present experienceswith chemical flower and fruit thinning of apple and pear, amplifiedwith some data of recent trials with known and new flower thinners(mainly ethephon and ammonium thiosulphate) and fruit thinners (mainly1-napthylacetic acid (NAA), its amide (NAAm), carbaryl, ethephon,conjugates of NAA and NAAm and benzyladenine). Chemical-thinningpractices give quite unpredictable results. This inconsistency is atleast partly caused by weather factors, such as temperature and airhumidity, but tree factors are also involved. To solve this problem,climate-room and multi-site trials are proposed, together with anintergrated approach in elucidating background mechanisms and thedevelopment of new fruit thinning compounds.     

Fruit size in a tropical tree species: variation,preference by birds,and heritability

Ocotea tenera (Lauraceae), an understory bird-dispersed tree, produces single-seeded fruits that vary in diameter from 1.4 to 2.4 cm. Much of the variation within a population at Monteverde, Costa Rica occurred within individual trees. The relative size of fruits produced by different trees remained generally constant over an 11-year period despite slight differences between years in the average size of fruits produced by a given tree.Fruit-eating birds could thus express their preferences for particular fruit size characteristics by choosing among trees that have distinct distributions of fruit diameters, and between individually variable fruits within trees. In a field study of individually marked fruits, birds removed 46.2% of fruits; the rest of the fruits were destroyed by invertebrate (25.3%) and vertebrate (4.3%) pulp-feeders or aborted by the plant after remaining ripe but uneaten for as long as 100 days (24.2%). The four major avian seed dispersers of O. tenera each have gape widths exceeding all but the largest fruits. Birds preferred plants with greater-than-average-sized fruits; within trees, they favored larger fruits, apparently because net pulp mass increases with fruit diameter. Fruits that ripened early in the season were more likely to be removed and were removed more quickly than late-ripening fruits.Based on mother-offspring regressions of mean fruit size, the phenotypic variation in fruit diameter in O. tenera is highly heritable, indicating the potential for an evolutionary response to selection by birds. Nonetheless, directional selection on fruit size or shape is likely to be inconsistent, constrained by genetic correlations, and weak compared to selection on traits like fecundity or phenology.