Phenology of Carpophilus spp. (Coleoptera: Nitidulidae)in stone fruit orchards as determined by pheromone trapping: Implications for prediction of crop damage

Traps baited with synthetic aggregation pheromone and fermenting bread dough were used to monitor seasonal incidence and abundance of the ripening fruit pests, Carpophilus hemipterus (L.), C. mutilatus Erichson and C. davidsoni Dobson in stone fruit orchards in the Leeton district of southern New South Wales during five seasons (1991-96). Adult beetles were trapped from September-May, but abundance varied considerably between years with the amount of rainfall in December-January having a major influence on population size and damage potential during the canning peach harvest (late February-March). Below average rainfall in December-January was associated with mean trap catches of < 10 beetles/trap/week in low dose pheromone traps during the harvest period in 1991/92 and 1993/94 and no reported damage to ripening fruit. Rainfall in December-January 1992/93 was more than double the average and mean trap catches ranged from 8-27 beetles/week during the harvest period with substantial damage to the peach crop. December-January rainfall was also above average in 1994/95 and 1995/96 and means of 50-300 beetles/trap/week were recorded in high dose pheromone traps during harvest periods. Carpophilus spp. caused economic damage to peach crops in both seasons. These data indicate that it may be possible to predict the likelihood of Carpophilus beetle damage to ripening stone fruit in inland areas of southern Australia, by routine pheromone-based monitoring of beetle populations and summer temperatures and rainfall.

Developing improved supply chains for temperate fruits in transitional Asian economies of Thailand and Vietnam.

Asia's increasing demand for both tropical and temperate fruit is projected to grow significantly. Compared with most developed countries, the production of temperate fruits (peach, nectarine, plum and apple) has expanded rapidly in China over the past 20 years. In contrast, current production of plums and peaches in neighbouring countries (Thailand and Vietnam) is very low but their fruit enters the market earlier. Thailand and Vietnam have enormous potential to satisfy a market window in the northern hemisphere period from March to May inclusive when there is little or no stone fruit on the Asian market. In Vietnam, fruit is harvested in an immature state to avoid disease and fruit fly problems and consequently lacks size and flavour. Approximately 30-40% of locally produced fruit in Vietnam does not reach market due to disease and poor handling during picking and transport. In Thailand, much of the infrastructure needed to transport, store, process and market temperate fruits successfully are now in place. However, there are currently no cool chain management or quality assurance systems to ensure a fresh product reaches the consumer with minimal deterioration. In Vietnam, growing stone fruit under the traditional system with little or minimal inputs, the farmer may receive between AUD3,000-5,000 per ha. In comparison, under higher input systems incorporating fertiliser, irrigation and pest and disease management, net returns can be increased seven-fold. Strengths and weaknesses of the current supply chains in these two countries are discussed.

Insecticide resistance and implications for future aphid management in Australian grains and pastures: A review

Aphids can cause substantial damage to cereals, oilseeds and legumes through direct feeding and through the transmission of plant pathogenic viruses. Aphid-resistant varieties are only available for a limited number of crops. In Australia, growers often use prophylactic sprays to control aphids, but this strategy can lead to non-target effects and the development of insecticide resistance. Insecticide resistance is a problem in one aphid pest of Australian grains in Australia, the green peach aphid (Myzus persicae). Molecular analyses of field-collected samples demonstrate that amplified E4 esterase resistance to organophosphate insecticides is widespread in Australian grains across Australia. Knockdown resistance to pyrethroids is less abundant, but has an increased frequency in areas with known frequent use of these insecticides. Modified acetylcholinesterase resistance to dimethyl carbamates, such as pirimicarb, has not been found in Australia, nor has resistance to imidacloprid. Australian grain growers should consider control options that are less likely to promote insecticide resistance, and have reduced impacts on natural enemies. Research is ongoing in Australia and overseas to provide new strategies for aphid management in the future.

Vermiculite improves early development and survival of low chill stone-fruit embryos rescued in vitro

There are many reports of efficient embryo germination and the method has been optimized to suit subtropical low chill genotypes. However the subsequent growth, vigor, and ability of germinated embryos to develop and survive acclimatization is rarely reported. Many germinated embryos do not survive acclimatization, develop slowly, or fail to develop normally. Methods to improve plant development from in vitro embryo cultures are needed to improve the number of plants that survive to be useful in breeding programs. This paper describes an improved method of embryo rescue that significantly increases embryo shoot and root development that leads to increased plant survival. Four treatments: Woody Plant Media (WPM) solidified with agar, vermiculite with liquid WPM, vermiculite with WPM plus agar, and conventional stratification, were evaluated for embryo growth and subsequent plantlet development and survival for two low-chill peach and one low-chill nectarine cultivar. Highly significant improvements were found for shoot and root development of seedlings germinated in vermiculite based media compared to embryos germinated in conventional agar-based media. Vermiculite with WPM and agar improved plantlet growth subsequent to in vitro culture and significantly increased survival of germinated embryos resulting in more plants reaching the field.

SF06016 Low Chill Peaches Online

The aim of this project is to bring information on low chill stonefruit varieties to a user in a clear and friendly format to aid in that decision process. Low Chill Australia see this project as high priority for its members to be competitive by growing high quality, early season peach and nectarine fruit varieties. Data will be collated from grower surveys, breeder’s descriptions and literature, and entered into an Access Database and published on the web for stonefruit growers in tropical and sub-tropical regions across Australia. Links will be available from the Low Chill Australia and Summerfruit Australia websites.

Preference and performance of Diachasmimorpha kraussii (Fullaway) (Hymenoptera: Braconidae) on five commercial fruit species

Diachasmimorpha kraussii is a polyphagous endoparasitoid of dacine fruit flies. The fruit fly hosts of D. krausii, in turn, attack a wide range of fruits and vegetables. The role that fruits play in host selection behaviour of D. kraussii has not been previously investigated. This study examines fruit preference of D. kraussii through a laboratory choice-test trial and field fruit sampling. In the laboratory trial, oviposition preference and offspring performance measures (sex ratio, developmental time, body length, hind tibial length) of D. kraussii were investigated with respect to five fruit species [Psidium guajava L. (guava), Prunis persica L. (peach), Malus domestica Borkh. (apple), Pyrus communis L. (pear) and Citrus sinensis L. (orange)], and two fruit fly species (Bactrocera jarvisi and B. tryoni). Diachasmimorpha kraussii responded to infested fruit of all fruit types in both choice and no-choice tests, but showed stronger preference for guava and peach in the choice tests irrespective of the species of fly larvae within the fruit. The wasp did not respond to uninfested fruit. The offspring performance measures differed in a non-consistent fashion between the fruit types, but generally wasp offspring performed better in guava, peach and orange. The offspring sex ratio, except for one fruit/fly combination (B. jarvisi in apple), was always female biased. The combined results suggest that of the five fruits tested, guava and peach are the best fruit substrates for D. krausii. Field sampling indicated a non-random use of available, fruit fly infested fruit by D. kraussii. Fruit fly maggots within two fruit species, Plachonia careya and Terminalia catappa, had disproportionately higher levels of D. krausii parasitism than would be expected based on the proportion of different infested fruit species sampled, or levels of fruit fly infestation within those fruit.

Evaluation of exclusion netting for insect pest control and fruit quality enhancement in tree crops

This work evaluated the following aspects of the use of exclusion netting in low chill stone fruit: the efficacy of protection from fruit fly for this highly susceptible crop; the effects on environmental factors; and the effects on crop development. Concurrently, an economic viability study on the use of exclusion netting was undertaken. The trial site was a 0.6-ha block of low chill stone fruit at Nambour, south-east Queensland, Australia. In this area, populations of Queensland fruit fly (Bactrocera tryoni) are known to be substantial, particularly in spring and summer. The trial block contained healthy 4-year-old trees as follows: 96 peach trees (Prunus persica cv. Flordaprince) and 80 nectarine trees (40 P. persica var. nucipersica cv. White Satin and 40 P. persica var. nucipersica cv. Sunwright). Exclusion netting was installed over approximately half of the block in february 2001. The net was a UV-stabilized structural knitted fabric made from high-density polyethylene yarn with a 10-year prorated UV degradation warranty. The results demonstrated the efficacy of exclusion netting in the control of fruit flies. Exclusion netting increased maximum temperatures by 4.4 deg C and decreased minimum temperatures by 0.5 deg C. Although exclusion netting reduced irradiance by approximately 20%, it enhanced fruit development by 7-10 days and improved fruit quality by increasing sugar concentration by 20-30% and colour intensity by 20%.

Innovative new production systems for low-chill stonefruit in Australia and South-East Asia: a review

Fruit size and quality are major problems in early-season stonefruit cultivars grown in Australia and South-East Asia. In Australia, Thailand and Vietnam, new training and trellising systems are being developed to improve yield and fruit quality. Australian trials found that new training systems, such as the Open Tatura system, are more productive compared with standard vase-trained trees. We established new crop-loading indices for low-chill stonefruit to provide a guide for optimum fruit thinning based on fruit number per canopy surface and butt cross sectional area. Best management practices were developed for low-chill stonefruit cultivation using growth retardants, optimizing leaf nitrogen concentrations and controlling rates and timing of irrigation. Regulated deficit irrigation (RDI) improved fruit sugar concentrations by restricting water application during stage II of fruit growth. New pest and disease control measures are also being developed using a new generation of fruit fly baits. Soft insecticides such as (Spinosad) are used at significantly lower concentrations and have lower mammalian toxicity than the organophosphates currently registered in Australia. In addition, fruit fly exclusion netting effectively eliminated fruit fly and many other insect pests from the orchard with no increase in diseases. This netting system increased sugar concentrations of peach and nectarine by as much as 30%. Economic analyses showed that the break-even point can be reduced from 12 to 6 years Open Tatura trellising and exclusion netting.

Effects of light conditions and plant density on growth and reproductive biology of Cascabela thevetia (L.) Lippold

Cascabela thevetia (L.) Lippold (Apocynaceae) is an invasive woody weed that has formed large infestations at several locations in northern Australia. Understanding the reproductive biology of C. thevetia is vital to its management. This paper reports results of a shade house experiment that determined the effects of light conditions (100% or 30% of natural light) and plant densities (one, two, four or eight plants per plot) on the growth, time to flowering and seed formation, and monthly pod production of two C. thevetia biotypes (peach and yellow). Shaded plants were significantly larger when they reached reproductive maturity than plants grown under natural light. However, plants grown under natural light flowered earlier (268 days compared with 369 days) and produced 488 more pods per pot (a 5-fold increase) over 3 years. The yellow biotype was slightly taller at reproductive maturity but significantly taller and with significantly greater aboveground biomass at the end of the study. Both biotypes flowered at a similar time under natural light and low plant densities but the yellow biotype was quicker to seed (478 versus 498 days), produced significantly more pods (364 versus 203 pods) and more shoot growth (577 g versus 550 g) than the peach biotype over 3 years. Higher densities of C. thevetia tended to significantly reduce the shoot and root growth by 981 g and 714 g per plant across all light conditions and biotypes over 3 years and increase the time taken to flower by 140 days and produce seeds by 184 days. For land managers trying to prevent establishment of C. thevetia or to control seedling regrowth once initial infestations have been treated, this study indicates that young plants have the potential to flower and produce seeds within 268 and 353 days, respectively. However, with plant growth and reproduction most likely to be slower under field conditions, annual surveillance and control activities should be sufficient to find and treat plants before they produce seeds and replenish soil seed banks. The most at-risk part of the landscape may be open areas that receive maximum sunlight, particularly within riparian habitats where plants would consistently have more favourable soil moisture conditions.

The growth, reproduction and survival of 'Cascabela thevetia' seedlings under two levels of canopy cover

The ornamental tree 'Cascabela thevetia', from tropical America, has naturalised and formed large infestations at several locations in northern Australia. Some understanding of its ecology and invasiveness was gleaned from a field experiment undertaken in North Queensland. The experiment quantified the growth, time to seed formation and survival of seedlings of the peach biotype growing under light and dense canopy cover within a riparian habitat. Growth, reproduction and survival of young plants varied. Growth was most rapid for seedlings away from, or on the edge of infestations because they were constrained by parent plants. The findings also suggested that land managers have at least 12 months following control to detect new plants, or regrowth, before plants set seed and replenish soil seed banks.

Dynamics of edge dislocations in a sheared lamellar mesophase

The dynamics and interactions of edge dislocations in a nearly aligned sheared lamellar mesophase is analysed to provide insights into the relationship between disorder and rheology. First, the mesoscale permeation and momentum equations for the displacement field in the presence of external forces are derived from the model H equations for the concentration and momentum field. The secondary flow generated due to the mean shear around an isolated defect is calculated, and the excess viscosity due to the presence of the defect is determined from the excess energy dissipation due to the secondary flow. The excess viscosity for an isolated defect is found to increase with system size in the cross-stream direction as L-3/2 for an isolated defect, though this divergence is cut-off due to interactions in a defect suspension. As the defects are sheared past each other due to the mean flow, the Peach-Koehler force due to elastic interaction between pairs of defects is found to cause no net displacement relative to each other as they approach from large separation to the distance of closest approach. The equivalent force due to viscous interactions is found to increase the separation for defects of opposite sign, and decrease the separation for defects of same sign. During defect interactions, we find that there is no buckling instability due to dilation of layers for systems of realistic size. However, there is another mechanism, which is the velocity difference generated across a slightly deformed bilayer due to the mean shear, which could result in the creation of new defects. (C) 2013 AIP Publishing LLC.

桃胶多糖的化学研究及其应用

毛桃(Prunus persica (L.) Batsch)是我国普遍裁培的一种果树。在桃树致病或受伤处会有桃胶(Peach gum)分泌。在一般情况下，天然桃胶是很难溶于水的，因此限制了它的使用。本文首次通过碱溶后，以快速沉淀的方法来降低桃胶的局部结晶度，使得桃胶能较顺利地溶解于冷水中。并提出了由于桃胶对金属镁的强络合作用而影响了桃树的正常生长这一推测。 桃胶经碱溶纯化制得钠型胶，再用阳离子交换树脂处理得氢型胶。用不同的方法处理桃胶后，桃胶红外光谱的结构特征吸收方才显示出来。钠型胶具840Cm-1的红外吸收;氢型胶具890Cm-1、1730cm-1的红外吸收;甲基化后的桃胶具810Cm-1、870Cm-1、890Cm-1的红外吸收。桃胶的比旋光度为负值。核磁共振证明桃胶多糖中，α-和β-甙键共存。碱溶纯化的桃胶多糖经凝胶色谱和超速离心证明为单一物质。 桃胶多糖不具还原性。经气相层析和比色法分析，桃胶中D-葡萄糖醛酸(包括4-甲氧基-D-葡萄糖醛酸）、D-半乳糖、D-木糖、D-甘露糖、L-阿拉伯糖、L-鼠李糖含量分别为10%、30%、11%、1.3%、45%、2%。18C-NMR和甲基化分析，证明桃胶大分子链是高度支链化的。且阿拉伯糖和半乳糖在大分子结构中有呋喃环和吡喃环两种形式共存。桃胶按物理性状，可分为两种。它们仅仅是在分子量和水溶液中的粘度不同，分子量( Mw)为342.9万和416.9万，特性粘度(η)为0.425dl/g和0.866dl/g。但在化学成分及结构特征上是相似的。以甘油为增塑剂，能增加桃胶成膜后的柔韧性。

采后果实衰老和冷害的机理及调控研究

衰老和冷害是果实采后极易发生的两种重要的生理性失调，严重影响了果实的商品质量和市场价值。本论文选用在采后贮藏期间容易衰老和出现冷害的枇杷、杧果和桃果实作为研究材料，比较深入系统地研究了这些果实采后衰老或冷害的机理和调控。主要的研究内容包括：（1）枇杷果实在不同贮藏条件下（低温、气调）的生理代谢、品质变化、衰老进程和贮藏性;（2）杧果果实采后生理特性及外源草酸和水杨酸处理对杧果冷害、品质和活性氧代谢的影响;（3）桃果实在低温贮藏下冷害发生特征，并从氧化还原平衡和蛋白质组表达方面探讨了桃果实采后冷害的发生机理。 试验结果表明：（1）与自发气调包装（MAP）相比，气调（CA, 10% O2＋1%CO2）能更有效的抑制枇杷果实的腐烂、延缓衰老、保持品质，以及控制多聚半乳糖醛酸酶（PG）和多酚氧化酶（PPO）的活性。枇杷果实在1 ºC气调下贮藏50天，腐烂指数低于7%，并保持固有的风味品质。短时间的高氧处理对果实腐烂和品质影响不大，促进了贮藏后期果实内乙醇的积累;（2）外源草酸和水杨酸处理显著减轻了杧果果实在贮藏期间的冷害程度，但对果实的硬度、可溶性固型物和可滴定酸含量影响不大。外源草酸和水杨酸提高了果实内抗坏血酸和谷胱甘肽的还原状态、提高了H2O2含量、降低了O2-含量和脂氧合酶活性，并提高了超氧化物岐化酶、过氧化氢酶、过氧化物酶、抗坏血酸氧化酶和谷胱甘肽还原酶的活性;（3）与非冷害桃果实相比，冷害桃果实呈现较低的H2O2含量、较高的膜透性、较高的酚类物质含量以及较高的抗坏血酸和谷胱甘肽还原状态。水杨酸对果实内H2O2含量、膜透性、酚类物质含量有显著的影响。冷害桃果实与非冷害桃果实的蛋白质表达也有明显的区别，32个差异表达蛋白依其功能分为8类，分别为能量产生、代谢、次生代谢、抗性、蛋白质的分布、基因转录、细胞结构和细胞生长。水杨酸对其中20个蛋白质的表达有显著影响。烯醇化酶、脂质运载蛋白、脱水蛋白、分支酸变位酶以及桂醇脱氢酶等与桃冷害发生有密切关系。 通过对上述结果的分析，得出如下结论：（1）与MAP相比，CA（10% O2 + 1% CO2）可显著延缓枇杷果实的衰老、腐烂和品质下降，CA延缓枇杷衰老的作用机理可能与其减轻果实内氧胁迫、降低PG和PPO活性有关;（2）用外源草酸或水杨酸处理杧果，可减轻贮藏期间果实的冷害程度，其调控机制可能与提高果实抗坏血酸和谷胱甘肽的还原状态、抑制O2－含量和提高H2O2含量有关;（3）在低温贮藏下，桃冷害发生与果实内较低的H2O2含量、生物膜完整性的破坏以及酚类物质含量增加密切相关，果实生物膜完整性的破坏与烯醇化酶、脂质运载蛋白、过敏蛋白以及脱水蛋白的下调表达有关，而酚类物质含量的增加与分支酸变位酶、桂醇脱氢酶的上调表达有关。

不同葡萄糖/果糖类型桃果实发育期间可溶性糖的变化规律和代谢

以成熟果实中不同葡萄糖/果糖（G/F）类型的6个桃品种（G/F≈1品种：‘冈山白’、‘山一白桃’和‘燕红’;高G/F品种：‘张黄7号’、‘龙246’和‘临白7号’）为试材，采用高效液相色谱法测定果实发育期果实和叶片中可溶性糖含量，并在盛花后74 d或101 d测定了‘冈山白’、‘山一白桃’、‘张黄 7号’和‘龙 246’新梢韧皮部中可溶性糖的含量;测定了果实发育过程中‘山一白桃’和‘临白7号’果实中的可溶性糖和淀粉代谢相关酶的活性。研究成熟果实中不同G/F类型桃果实内G/F差异的部位和时期;分析桃果实内G/F差异的可溶性糖代谢调控机理。 成熟果实中不同G/F类型桃果实中均以蔗糖作为主要碳水化合物积累形式，花后43–85 d蔗糖含量很低，随后持续快速积累直至果实成熟;花后43–85 d山梨醇有升高趋势，在果实成熟前40 d左右迅速降低;葡萄糖和果糖含量在果实发育早期较高，之后逐渐降低;但两类不同G/F桃在整个果实发育过程中G/F值与果实成熟时相似。叶片中贮藏的可溶性糖主要是蔗糖和山梨醇，在果实整个发育期间，G/F≈1品种叶片中G/F约1-3，而高G/F品种叶片中G/F约为2-7。G/F≈1品种‘冈山白’和‘山一白桃’与高G/F品种‘张黄 7号’和‘龙 246’韧皮部中山梨醇占总可溶性糖47-63％，显著高于蔗糖、葡萄糖和果糖的含量，G/F为0.8-0.91，且两类不同G/F桃品种间G/F值不存在显著差异。 成熟果实中G/F≈1类型的‘山一白桃’和高G/F值类型的‘临白7号’整个果实发育过程中，葡萄糖、山梨醇和淀粉的含量在这两个品种间一般没有明显差异;‘山一白桃’果实中的果糖含量显著高于‘临白7号’果实中的果糖;果实最后迅速生长期，‘山一白桃’果实中的蔗糖明显高于‘临白7号’。‘山一白桃’和‘临白7号’果实中的NAD+依赖型山梨醇脱氢酶（NAD+-SDH）活性低，两者有相似的变化趋势，一般无显著差异。‘临白7号’果实中的NADP+依赖型山梨醇脱氢酶（NADP+-SDH）和山梨醇氧化酶（SOX）活性一直高于‘山一白桃’，两者NADP+-SDH和SOX的活性分别在花后93-123 d和花后43-93 d有显著差异。‘临白7号’果实中的果糖激酶（FK）活性一般高于‘山一白桃’。花后43-93 d，‘临白7号’果实中的磷酸蔗糖合成酶（SPS）和蔗糖合成酶（SS）活性一般显著‘山一白桃’。果实最后迅速生长期，蔗糖快速积累，葡萄糖、果糖、山梨醇和淀粉含量迅速降低，同时伴随有SPS和SS活性的迅速升高。在整个果实发育过程中，两个品种果实中的淀粉酶活性较高，其果实中的淀粉含量和淀粉酶活性都有明显的下降趋势。 研究结果表明，整个果实发育过程中桃果实中均存在G/F≈1和高G/F现象，光合产物在韧皮部的运输对桃果实的G/F没有显著的影响，果实中G/F的差异主要由于果实内糖代谢差异所导致。‘临白7号’果实中山梨醇向果糖方向的转化能力与‘山一白桃’一般没有显著差异，由于不同时期较高的NADP+-SDH和SOX活性，使得山梨醇向葡萄糖方向的转化能力明显高于‘山一白桃’，同时，‘临白7号’果实中的FK活性一般高于‘山一白桃’，因此导致‘临白7号’果实中G/F高于‘山一白桃’。

桃和苹果库源关系中源叶光合作用，光抑制及其光保护机制的研究

以‘早久保’（Prunus persica (L.) Batch.）为试材，在果实最后迅速生长期，通过去果处理降低库力，同时设留果对照，并通过环剥和保留相同数量叶片严格控制库源关系，进行了源叶净光合速率（Pn）、叶绿素荧光、叶黄素循环、抗氧化酶及抗氧化同化物日变化的研究。结果表明，和留果对照相比，去果处理显著降低了源叶Pn、气孔导度（gs）和蒸腾速率（E），但显著增加了胞间二氧化碳浓度（Ci）、叶面饱和蒸汽压亏缺（VPDl）和叶片温度（Tl）。光系统II光化学效率（ΦPSII）以及羧化速率（CE）与Pn平行降低。中午去果降低Pn主要归因于非气孔限制。在低库需条件下，开放的PSII反应中心捕获能量的降低以及关闭的PSII反应中心的增加导致了ΦPSII的降低。去果处理叶片中依赖于叶黄素循环的热耗散以及抗氧化系统的上调保护叶片免受光氧化破坏。和留果对照相比，去果处理的叶片有更大的叶黄素循环库，更高的脱环氧化状态以及更高的抗氧化酶活性，包括超氧化物歧化酶（SOD）、抗坏血酸过氧化物酶（APX）、单脱氢抗坏血酸还原酶（MDAR）和脱氢抗坏血酸还原酶（DHAR）的活性以及更高的还原型抗坏血酸（AsA）和还原型谷胱甘肽（GSH）的含量。但与此同时，去果显著增加了过氧化氢（H2O2）以及丙二醛（MDA）的含量，这意味着在去果处理的叶片中可能会发生光氧化破坏。 以一年生‘皇家嘎拉’苹果（Malus domestica Borkh.）组培苗为试材，通过环剥降低库力，进行了源叶Pn、叶绿素荧光、核酮糖-1,5-二磷酸羧化酶/氧化酶（Rubisco）以及光系统II（PSII）复合体关键蛋白PsbA和PsbO含量日变化的研究。和对照相比，环剥显著降低了源叶Pn、gs和E，但是却显著增加了Ci、Tl和淀粉的含量。在低库需下，开放的PSII反应中心捕获能量的降低以及关闭的PSII反应中心的增加导致了ΦPSII的降低。另一方面，环剥降低了光合作用关键酶Rubisco以及PSII复合体PsbA和放氧复合体PsbO的含量。以上结果表明，环剥降低Pn主要归因于非气孔限制。