苜蓿轮作玉米后土壤养分时空变化特征分析

以7龄紫花苜蓿地为前茬,连续3年采用同一高产栽培方案种植玉米,研究其土壤养分的时空变化特征。结果表明,苜蓿轮作玉米后,随着玉米种植年限的增加,玉米各生育时期土壤有机质、全氮和全磷含量均有不同程度下降。其中,0-20cm土层降幅高于其他土层,且降幅表现为有机质>全氮>全磷。受玉米高量施氮的影响,碱解氮含量随玉米种植年限的增加而增加,不同生育时期、不同土层年际间碱解氮含量的差异多数达到显著水平。玉米产量随着连作年限的增加而降低,第2,3年产量较第1年产量分别下降了9.7%和13.6%,表明苜蓿对后作的增产效应持续降低。     

不同施氮量对夏玉米产量、氮肥利用率及氮平衡的影响

通过田间小区试验研究了不同施氮量对夏玉米产量、氮肥利用率、硝酸盐淋溶及氮平衡的影响。结果表明，施氮对夏玉米子粒有显著的增产作用，但随施氮量的增加产量变化不大。氮肥利用率在9．2％-22．6％之间，随施氮量的增加而降低。施氮可明显提高0-160cm剖面土壤NO3^--N含量，而且随深度的增加NO3^--N含量呈降低趋势，累积峰主要在20—60cm之间。玉米收获后，随着施氮量的增加氮素的损失量增加，各施氮处理的硝态氮残留量在121～221kg／hm^2之间，以N250处理的残留量最高，残留率近65％。     

不同类型氮肥对夏玉米产量、氮肥利用率及土壤氮素表观盈亏的影响

通过田间小区试验,比较研究了不同肥料类型、不同施氮量对夏玉米产量、氮肥利用率、土壤氮素表观盈亏量及植株氮素累积萤的影响.结果表明,夏玉米产量随施氮量增大而增大,但施肥过量产量降低,同等施氮条件下,控释掺混氮肥各处理夏玉米产量和氮肥利用率均高于普通尿素处理.施氮显著增加了0-40cm土层土壤无机氮含量,同时显著提高了玉米生长中后期植株体内氮素含量.经夏玉米季后,除高量施氮的T4,T6处理出现土壤氮索表观盈余外,其余处理均出现土壤氮素表观亏缺,施氮后土壤表观盈余量增加,且随施氮量的增加而增加.     

不同培肥措施对银北灌区土壤盐碱特性、玉米生长及产量指标的影响

为探究银北灌区盐碱地不同培肥措施下的土壤盐碱特性和作物生长情况，通过3年定位试验开展了不同量化肥、秸秆、生物有机肥应用后对土壤盐碱特性、春玉米生长及产量指标的影响研究。结果表明:(1)玉米播前土壤表现为对照脱盐碱化现象最明显，而高量生物有机肥和高量化肥有增加耕层土壤盐分含量的趋势，秸秆还田在降低玉米生长前期耕层土壤pH值方面较显著；(2)高量秸秆还田在提高玉米株高和增加玉米茎粗方面优于施用生物有机肥和中低量化肥，玉米株高、茎粗随着有机肥和秸秆用量的增加而提高；(3)在增加玉米穗长、穗粗和穗粒重方面，单施化肥措施优于单施秸秆或生物有机肥；在增加玉米百粒重方面，秸秆还田与化肥措施效果相当；培肥第3年各处理玉米产量表现为:施用生物有机肥T4>T3>T2，施用秸秆T6>T7=T5，施用化肥T10>T9>T8，配施处理T11>T12，玉米产量随生物有机肥和化肥施用量的增加而提高，秸秆还田在提高玉米产量方面优于生物有机肥，而中量秸秆还田在提高玉米产量方面优于高量和低量秸秆还田。与施用化肥相比，试验区连续3年配施生物有机肥在提高玉米产量方面已表现出优势，而配施秸秆在提高玉米产量方面尚未发挥优势，秸秆还田相对生物有机肥在提高作物产量方面存在滞后效应。     

磷酸胆碱合剂对不同种植密度玉米叶片衰老生理的影响

针对东北地区低温冷凉气候区密植条件下玉米叶片早衰现象,于大田条件下,研究了不同种植密度下磷酸胆碱合剂对京单28和先玉335叶片衰老相关指标及产量性状的影响。结果表明,随种植密度增加,玉米叶片保绿度、叶绿素含量下降;超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)活性降低,丙二醛(MDA)含量升高。化控处理后,不同种植密度下,2个玉米品种叶绿素含量提高了6.57%~23.40%,保绿度提高了4.5%~13.9%;SOD、POD、CAT活性在中高密度条件下较对照升高了4.76%~21.64%,MDA含量较对照降低4.39%~11.82%;穗长、行粒数、百粒重和产量均较对照有所增加,京单28产量最高增加了398.3kg/hm2,先玉335产量最高增加了774.2kg/hm2。以不同品种不同密度下的各生理指标与产量进行相关分析,结果表明除MDA含量与产量呈极显著负相关外(r=-0.810**),其余都表现出正相关,其中POD活性(r=0.660*)达到了显著水平。表明磷酸胆碱合剂可以延缓玉米叶片的衰老。     

连年深松和施用有机肥对土壤肥力及玉米产量的影响

通过8年的田间定位试验,研究了不同耕作方法及有机肥用量对春玉米产量和土壤理化性状的影响。结果表明,连年深松能显著降低10~20 cm和20~30 cm土壤体积质量,施用有机肥30 000 kg/hm~2以上能显著降低10~20 cm土壤体积质量;深松导致耕层土壤有机质、全氮和全磷含量呈下降趋势,碱解氮和速效磷略有增加;耕层土壤有机质、全氮、碱解氮和速效磷含量随有机肥用量的增加而显著增加,全磷、全钾和速效钾含量增加幅度与有机肥用量无关;深松和施用有机肥均能显著增加玉米产量,施用有机肥的增产效果好于深松,但其增产速率随有机肥用量增加而降低。连年深松对改善土壤理化性质和提高玉米产量的效果好于隔年深松,且在施用有机肥的条件下效果更好。     

氮、磷营养对黄土高原旱地玉米产量、品质的影响机理研究

田间裂区试验研究了不同氮磷施用水平对旱地玉米子粒产量和品质的影响。结果表明,增施氮、磷肥,能增加子粒粗蛋白、粗脂肪、赖氨酸含量和子粒容重,降低粗淀粉的含量,提高玉米产量和水分利用效率,且氮肥效果优于磷肥。施用氮、磷肥能够增加土壤碱解氮、速效磷的浓度,有利于玉米根系对氮磷营养的吸收;在促进根系生长的同时,提高玉米的生物产量,增加植株茎秆、叶片对氮素、磷素的吸收、转化以及向穗轴、子粒的输送,显著提高子粒的灌浆强度,促进灌浆中后期子粒氮、磷的积累,增大粒重,改善子粒品质。     

高磷土壤施用锌肥对玉米氮磷钾吸收和积累的影响

在田间高磷土壤条件下,研究不同锌肥施用量对玉米各个生育期植株氮磷钾吸收和积累以及对产量的影响,以期为提高氮磷钾的利用率、玉米增产和合理施锌肥提供科学依据。研究结果表明:施用锌肥能提高玉米各生育期植株和籽粒N和K的含量,P的含量则呈降低的趋势。同一施锌量,玉米植株积累的NPK量随植株生长而增加,在同一生育期中,玉米植株积累的N和K量在施锌量10～50 kg/hm2范围内,呈增加的趋势,继续增加施锌量则呈降低的趋势,累积的P量在施锌量10～150 kg/hm2范围内呈降低的趋势,说明施锌量10～150 kg/hm2范围内,能够提高NK肥的利用率。不同施锌量对玉米产量的影响是施锌量10～50 kg/hm2范围内,呈增加的趋势,继续增加施锌量则呈降低的趋势。     

生长调节剂对玉米氮代谢的影响

采用田间小区试验,研究了植物生长调节剂不同配方对玉米光合特性、氮素代谢以及玉米子粒产量与蛋白质含量的影响。结果表明,调节剂不同配方处理能明显提高子粒灌浆期玉米功能叶叶绿素的含量和净光合速率,功能叶与茎秆中的全氮和蛋白氮的减少量,功能叶的硝酸还原酶、谷氨酰胺合成酶与转化酶活性及子粒谷氨酰胺合成酶与转化酶活性;灌浆中、后期玉米功能叶蛋白水解酶活性和玉米子粒产量、子粒蛋白质含量及蛋白质产量。在生长调节剂中,PGR4处理玉米子粒的蛋白氮含量比对照提高1.35.g/kg,子粒产量比对照增加9.5%,子粒蛋白质产量比对照增加19.6%;PGR3处理玉米子粒的蛋白氮含量比对照提高0.78.g/kg,子粒产量比对照增加11.4%,子粒蛋白质产量比对照增加17.4%,是较合理的配方。     

海藻酸复混肥料对玉米产量及品质的影响

为探讨不同海藻酸含量复混肥料对玉米产量及品质的影响,采用田间试验,以施基肥的方式,研究不同施肥处理条件下玉米品质及产量的变化。结果表明,施肥量相同的条件下,海藻酸复混肥料能显著增加玉米茎粗、穗长、百粒重,增加玉米籽粒、茎叶产量及籽粒中淀粉、蛋白质的含量,缩短玉米秃尖,提高玉米籽粒对氮、磷、钾养分的吸收,但对玉米株高影响不显著。     

增强紫外线-B辐射冬小麦产量和光合特性的影响

为了研究紫外线-B（ultraviolet-B UV-B, 280-320nm）辐射增强20%对保护性耕作冬小麦产量及光合特性的影响,在中国南京开展了2 a的田间试验研究。试验采用常规耕作（耕深25 cm）、少耕（耕深10 cm）和免耕（土壤不耕作）3种耕作处理种植冬小麦,采用人工增加紫外辐射的方法模拟UV-B辐射增强。试验测定了冬小麦旗叶的光合速率、光合-光响应曲线、叶绿素质量分数、超氧化物歧化酶（superoxide dismutase, SOD）活性、丙二醛（malonaldehyde, MDA）质量分数和可溶性蛋白质量分数等指标。结果表明：在UV-B辐射增强20%条件下,与常规耕作相比,少耕和免耕2种保护性耕作处理可显著提高冬小麦的产量,分别比常规耕处理作高1632.12和952.15 kg/hm2。从叶片光合生理特性来看,在UV-B辐射增强20%条件下,从冬小麦籽粒灌浆中期开始,少耕和免耕处理旗叶的光合速率、表观量子效率（apparent quantum yield, AQY）、最大净光合速率（Pmax）、叶绿素质量分数、SOD活性、可溶性蛋白质的质量分数显著高于常规耕作处理,MDA质量分数显著低于常规耕作处理,2种处理冬小麦的叶片衰老显著低于常规耕作处理。在UV-B辐射增强20%条件下,少耕和免耕2种保护性耕作处理冬小麦旗叶在籽粒灌浆中期及以后保持高的光合能力和低的衰老程度,是其产量高于常规耕作处理的原因。该文可为制定UV-B辐射增强条件下的冬小麦栽培措施提供参考。     

UV-B辐射增强对大麦生理生态的影响

在大田条件下,研究了UV-B辐射增强对大麦的生长发育、光合作用、蒸腾作用及其产量构成的影响。结果表明,UV-B辐射增强明显抑制大麦生长,使株高变矮、绿叶数减少、叶面积和干物质量下降,但抑制程度随生育期而异。在UV-B辐射增强条件下,大麦叶片叶绿素含量、净光合速率、气孔导度和蒸腾速率都有不同程度的降低,水分利用率也随之降低。UV-B辐射增强对大麦形态学和生理学上的不利影响,导致了大麦产量下降24.96%。     

全生育期UV-B辐射增强对棉花生长及光合作用的影响

植物光合系统是UV-B辐射最初和最重要的作用靶标。本文在大田条件下进行紫外灯照射处理,研究全生育期UV-B辐射增强(高于环境20%和40%)对棉花形态、干物质积累、光合色素和产量的影响,并通过分析棉花主茎功能叶片的气体交换参数和叶绿素荧光参数,探讨UV-B辐射增强影响棉花光合作用的机制。结果表明,UV-B辐射增强抑制了棉花生长和干物质积累,籽棉产量显著降低,且UV-B辐射越强,抑制作用越明显。随UV-B辐射的增强,棉花主茎功能叶的净光合速率(P_n)在各生育期均显著降低,叶绿素含量呈先升高后降低趋势,气孔导度(Gs)和蒸腾速率(Tr)未发生变化,胞间CO_2浓度(Ci)反而升高,说明P_n下降主要由非气孔限制因素造成。对叶绿素荧光参数的分析表明,PSⅡ的最大光化学量子产率(F_v/F_m)、实际光化学量子效率(ΦPSII)、线性电子传递速率(ETR)和光化学淬灭系数(qP)随着UV-B辐射的增强而降低,非光化学猝灭系数(NPQ)则显著升高,且各叶绿素荧光参数与Pn变化均显著相关;慢速弛豫NPQ(NPQS)及其在NPQ中的比例均随UV-B辐射的增强而显著提高,表明PSⅡ反应中心受损,光化学效率降低。以上结果证明,全生育期UV-B辐射增强降低了棉花的光合叶面积、叶绿素含量和净光合速率,引起棉花生长与物质积累受抑,产量降低。UV-B辐射增强引起的光合速率下降与PSⅡ反应中心遭到破坏密切相关。     

Field crop responses to ultraviolet-B radiation: a review

This paper provides an overview of existing literature on the ultraviolet-B (UV-B) radiation effects on field crops. Earlier reviews on field crop responses to UV-B considered few physiological processes or crops. For this review, we easily located about 129 studies on 35 crop species published since 1975. Here, we report the effects of UV-B radiation on visual symptoms, leaf ultrastructure and anatomy, photosynthetic pigments, UV-B absorbing compounds, photosynthesis, growth and development, yield, genotypic differences, and finally, interactions of UV-B with abiotic and biotic factors of crop plants. Experiments conducted in glasshouses, in closed and open top chambers, and under field conditions, with varying source (solar or artificial) and intensity of photosynthetically active radiation (PAR, 50–1800 μmol m⁻² s⁻¹) and UV-B (0–50 kJ m⁻² per day) are included. It is easy to conduct experiments that purport to evaluate the effects of projected UV-B intensities on crop species by providing supplemental irradiance with lamps or by reducing UV-B with filters; however, it is very difficult to simulate UV-B irradiance spectral changes that are likely to occur in nature. Collated results for each process are presented as percent change from control along with the experimental conditions in tabular format. Many of the studies showed dramatic effects of UV-B radiation, but under conditions with supplemental UV-B irradiance that was higher than would ever occur outside experimental conditions or in which the longer wavelengths in the PAR and UV-A, which moderate UV-B effects, were greatly reduced. Only 25 of the studies reviewed used experimental conditions and supplemental UV-B irradiance that approached realism. However, unrealistic the experimental conditions might be, an increase in understanding of basic plant physiology was gained from most of the studies.Visual symptoms consisting of chlorotic or necrotic patches on leaves exposed to UV-B were not unique. Both vegetative and reproductive morphology were altered by UV-B radiation. Leaf anatomy was altered due to changes in thickness of epidermal, palisade, and mesophyll layers. Enhanced UV-B generally decreased chlorophyll content (10–70%), whereas it increased UV-B absorbing compounds (10–300%) in many crops. Decrease in photosynthesis (3–90%), particularly at higher UV-B doses, was due to both direct (effect on photosystem) and indirect (decrease in pigments and leaf area) effects. The decreases in chlorophyll pigments and photosynthesis resulted in lower biomass and yield of most crop plants. Genotypes of crop species exhibited variability in leaf wax layer thickness, loss of chlorophyll, and increase in phenolics as mechanisms of tolerance to enhanced UV-B radiation resulting in changes in biomass/yield. Results from the few studies on interaction of UV-B with other abiotic and biotic factors did not lead to useful conclusions. Studies are needed to quantify the effects of UV-B radiation on crops in order to develop dose response functions that can facilitate development of dynamic simulation models for use in UV-B and other environmental impact assessments.     

Cotton responses to ultraviolet-B radiation: experimentation and algorithm development

The potential impact of an increase in solar ultraviolet-B (UV-B) radiation due to human activity on higher plants has been the subject of many studies. Little work has been carried out so far on cotton responses to enhanced UV-B radiation. The objective of this study was to determine whether or not the current and projected increases in UV-B levels affect cotton growth and development, and to quantify and develop UV-B radiation functional algorithms that can be used in simulation models. Two experiments were conducted during the summer of 2001 using sunlit plant growth chambers in a wide range of UV-B radiations under optimal growing conditions. Leaves exposed to UV-B radiation developed chlorotic and necrotic patches depending on the intensity and length of exposure. Along with changes in visible morphology, cotton canopy photosynthesis declined with increased UV-B radiation. The decline in canopy photosynthesis was partly due to loss of photosynthetic pigments and UV-B-induced decay of leaf-level photosynthetic efficiency (maximum photosynthesis) and capacity (quantum yield) as the leaves aged. The total leaf area was less due to smaller leaves and fewer leaves per plant. Less plant height was closely related to a shorter average internode length rather than a fewer mainstem nodes. The UV-B did not affect cotton major developmental events such as time taken to square, time to flower, and leaf addition rates on the mainstem. Lower biomass was closely related to both smaller leaf area and lower photosynthesis. The critical limit, defined as 90% of optimum or the control, for stem elongation was lower (8.7 kJ m⁻² per day UV-B) than the critical limit for leaf expansion (11.2 kJ m⁻² per day UV-B), indicating that stem elongation was more sensitive to UV-B than leaf expansion. The critical limits for canopy photosynthesis and total dry weight were 7 and 7.3 kJ m⁻² per day, respectively. The identified UV-B-specific indices for stem and leaf growth and photosynthesis parameters may be incorporated into cotton simulation models such as GOSSYM to predict yields under present and future climatic conditions.     

不同施肥处理对夏玉米产量及活性氧代谢的影响

以2009年建立的肥料定位试验为研究平台,于2013年进行了不同施肥处理对夏玉米产量及活性氧代谢影响的研究。结果表明,氮磷钾均衡施肥对夏玉米籽粒产量有显著的增产作用,产量达到9 299.85 kg/hm2,NPK处理的穗粒数和千粒重最高,NP处理的穗长、穗重与NPK处理没有显著差异;氮能有效地提高玉米穗位叶叶绿素含量和籽粒的灌浆速率;氮对玉米穗位叶过氧化物酶活性的影响最大,氮钾配施能够有效地提高玉米穗位叶超氧化物歧化酶活性,增加穗位叶可溶性蛋白质含量,降低穗位叶丙二醛含量。合理配施氮钾能够有效地提高玉米穗位叶光合效率,延缓叶片衰老,提高玉米籽粒产量。     

Elevated UV-B radiation modifies the extractability of carbohydrates from leaf litter of Quercus robur

We examined the influence of elevated UV-B radiation on the extractability of carbohydrates from leaf litter of Quercus robur. Saplings were exposed to a 30% elevation above the ambient level of erythemally weighted UV-B (280-315 nm) radiation for eight months at an outdoor facility. UV-B radiation was applied under arrays of fluorescent lamps filtered with cellulose diacetate, which transmitted both UV-B and UV-A (315-400 nm) radiation. Saplings were also exposed to elevated UV-A radiation under arrays of polyester-filtered lamps and to ambient radiation under arrays of non-energised lamps. Abscised leaves were collected, ground and sequentially treated with seven solvents in order to fractionate extractable carbohydrates based on the way in which they are held in the cell wall. Elevated UV-B radiation reduced the extractability of carbohydrates from cell walls of Q. robur. Sodium phosphate buffer at pH 7 extracted 10% less total carbohydrate from leaf material exposed during growth to elevated UV-B radiation under cellulose diacetate-filtered lamps than from leaf material grown under polyester-filtered and non-energised lamps. The cumulative amount of carbohydrate released by sequential extraction with phosphate buffer, CDTA, urea and sodium carbonate was between 5.1% and 7.8% lower from leaf material grown under cellulose diacetate-filtered lamps relative to that from leaves grown under non-energised lamps. Abscised leaves were also digested with Driselase, an enzyme mixture extracted from a basidiomycete fungus. No effects of elevated UV radiation were recorded on the amount of carbohydrate released by Driselase digestion. Regression analyses, using data from a previous field decomposition study, suggested that reduced availability of carbohydrates enhanced the colonisation of Q. robur litter by basidiomycete fungi, which then accelerated the decomposition rate of the litter in soil. We recommend that future studies into the effects of UV-B radiation on plant litter decomposition measure not only the concentrations of chemical constituents of litter, but also determine the availability of litter carbon sources to soil microbes, using methods similar to those used here.     

UV-B对人工气候室内黄瓜苗期生长、生理及抗氧化系统的影响

紫外线(UV-B)是植物生长发育的关键信号因子.过量或缺少UV-B都会影响作物的抗性、产量和品质.然而,目前植物工厂中适宜黄瓜生长的UV-B强度尚不明确.以黄瓜(Cucumis sativus L.)苗期植株为材料,研究不同强度UV-B对人工气候室内黄瓜苗期植株生长、生理和抗氧化系统的影响.结果表明:与对照相比,UV-B处理黄瓜植株高度降低4.2%~32.0%,叶片中可溶性蛋白含量降低14.2%~28.2%.3.33μmol/(m2·s)UV-B处理植株茎粗增加13.6%~22.3%,叶片中可溶性糖的含量增加22.7%~56.7%,同时激活抗氧化系统,超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)活性分别提高16.9%~23.2%,23.8%~25.9%,34.1%~50.4%,抗坏血酸含量增加27.4%~36.4%.由此可知,3.33μmol/(m2·s)UV-B有利于人工气候室中黄瓜苗期植株的生长发育、抗氧化酶活性提高及抗氧化物质生成.     

Ultraviolet-B radiation in a row-crop canopy: an extended 1-D model

A decrease in stratospheric ozone may result in a serious threat to plants, since biologically active short-wavelength ultraviolet-B (UV-B 280–320 nm) radiation will increase even with a relatively small decrease in ozone. Numerous investigations have demonstrated that the effect of UV-B enhancements on plants includes reduction in grain yield, alteration in species competition, susceptibility to disease, and changes in plant structure and pigmentation. To determine the physiological effects on plants of any increases in UV-B radiation, the irradiances at the potential sensitive plant surface need to be known. A number of radiative transfer models exist but because of the importance of sky diffuse radiation to the global UV-B irradiance, models designed to estimate photosynthetically active radiation or total solar radiation may not accurately model the UV-B. This paper compares spatially and temporally averaged measurements of the UV-B canopy transmittance of a relatively dense maize canopy (sky view: 0.27°) to the estimations of two one-dimensional models differing mainly in the handling of sky radiance. The model that considered the distribution of sky radiance tended to underestimate the canopy transmittance, the model that assumed an isotropic sky radiance distribution tended to overestimate the canopy transmittance. However, the assumption concerning the sky radiance distribution accounted for only about 0.01 of the model error. Consequently, the sky radiance distribution is probably not important in modeling such dense crop canopies. The model that overestimated transmittance and had the generally larger errors, a modified Meyers model, used the assumption of uniform leaf angle distribution, whereas in the other model, designated the UVRT model, leaf angle distributions were estimated by sample measurements. Generally this model would be satisfactory in describing the statistically average UV-B irradiance conditions in the canopy. This model may also be applied to other dense plant canopies including forests.     

外源水杨酸对UV-B增强下花生叶片光合特性的影响

通过大田模拟试验,设置自然光（CK）和UV-B辐射增强（E,增加量相当于当地4-5月UV-B辐射量的20%）两个大区,在每个大区内又分2个小区,从花针期（7月25日）开始,分别用蒸馏水（S0处理）和水杨酸水溶液（SA处理）连续3d在固定时段喷施花生植株的所有叶片后,用Li-6400型便携式光合作用测定仪观测和计算指定叶片的净光合速率（Pn）、蒸腾速率（Tr）、气孔导度（Gs）、胞间CO2浓度（Ci）和水分利用效率（WUE）的日变化过程,并进行对比分析。结果表明：不施水杨酸条件下（S0）,UV-B增强处理的花生叶片Pn、Tr、Gs和WUE的日均值比自然光（CK）处理分别下降35.7%、25.0%、25.0%、10.0%;而喷施水杨酸条件下（SA）则分别下降30.4%、17.9%、33.3%、19.4%,说明UV-B增强可降低花生叶片的净光合速率、蒸腾速率、气孔导度和水分利用效率,而喷施水杨酸可缓解UV-B增强对花生净光合速率的抑制作用,但并不能缓解UV-B增强对花生蒸腾作用、气孔导度及水分利用效率的抑制。