入侵豚草与本地植物竞争对丛枝菌根真菌多样性的影响

外来植物入侵常引起入侵地丛枝菌根真菌（arbuscular mycorrhizal fungi,AMF）多样性的变化,但AMF多样性的变化与入侵域本地植物种类相关性目前尚不清楚。本研究采用野外同质园试验研究豚草与不同本地植物竞争时对AMF多样性的影响。本试验共设入侵豚草单种、本地植物狗尾草Setaria viridis单种、藜 Chenopodium album单种、黄香草木犀Melilotus officinalis单种、豚草和狗尾草混种、豚草和藜混种、豚草和黄香草木犀混种以及裸土（无任何植物生长）对照8个处理。采集根区土样后分离、鉴定AMF,分析植株生物量和AMF物种多样性变化间的关系。结果表明,对豚草而言,单种处理中生物量均高于混种处理;豚草分别与3种本地植物竞争时,与黄香草木犀混种中豚草竞争能力最强,而与藜混种中竞争能力最弱。豚草分别与3种本地植物混种时土壤中AMF种丰度和香农-威纳指数分别显著高于豚草单种和本地植物单种,其中豚草与藜混种中AMF的种丰度和香农-威纳指数最高,与狗尾草混种中AMF的种丰度和香农-威纳指数最低,而豚草分别与3种本地植物混种处理中孢子密度显著低于本地植物单种（豚草与黄香草木犀混种除外）。豚草单种、狗尾草单种、藜单种、豚草分别与狗尾草和藜混种4个处理中AMF优势种均为网状球囊霉Glomus reticulatum和缩隔球囊霉Septoglomus constrictum;黄香草木犀单种中优势种是网状球囊霉、缩隔球囊霉和根内根孢囊霉Rhizophagus Intraradices,豚草与黄香草木犀混种中优势种是网状球囊霉和根内根孢囊霉;狗尾草单种、豚草与狗尾草混种和豚草与藜混种3个处理的特有种分别是木薯根孢囊霉Rhizoglomus manihotis、近明球囊霉Claroideoglomus claroideum和美丽盾巨孢囊霉Scutellospora calospora。狗尾草单种生物量与孢子密度呈显著正相关,豚草与狗尾草混种中豚草的生物量与均匀度呈显著负相关,豚草与黄香草木犀混种中豚草的生物量与孢子密度呈显著正相关,与香农威纳指数呈显著负相关。综上可知,豚草与本地植物竞争对AMF多样性的影响具有异质性,这与本地植物种类密切相关。     

氮水平和竞争对互花米草与芦苇叶特征的影响

 互花米草 (Spartina alterniflora) 和芦苇 (Phragmites australis) 是滨海盐沼湿地的多年生草本植物,从世界范围来看,它们二者具有区域性的相互入侵特征,因此研究生境条件对两物种互侵机制的影响是一个十分有意义的生态学命题。该文运用随机区组实验设计方法,模拟海滩环境、构建人工种群、控制可变因子,研究了外来种互花米草与本地种芦苇分别单种和混种时,叶特征对不同氮水平、不同植株密度的响应。结果表明：随着氮水平的升高,互花米草和芦苇的叶面积无论是在单种还是混种情况下都显著增加 (p<0.05),但混种条件下芦苇的叶面积在高氮水平下增幅减少,这与高氮状况下互花米草与芦苇的竞争加剧有关;氮水平对单种中两种植物的叶数影响最显著 (p<0.01),对混种中互花米草的叶数和芦苇的叶宽影响最大 (p<0.05)。植株密度增加导致种内和种间竞争加剧,无论在单种还是混种处理下,都造成两种植物叶面积的显著减少 (p<0.05)。单种处理中, 两物种的叶数受密度的响应最显著 (p<0.05);而混种处理中芦苇对互花米草的竞争显著减小了互花米草的叶宽和叶数(p<0.05),互花米草对芦苇的竞争则显著减小了芦苇 的叶长、叶宽和叶数 (p<0.05)。两种植物的竞争结果受到氮营养的调控,低、高氮水平下互花米草的种间竞争能力大于芦苇,中氮水平下则是芦苇的种间竞争能力大于互花米草。高氮水平下互花米草通过叶面积的快速增加抑制了芦苇的叶生长,使其叶面积减少,从而在竞争中占据优势,这可能是互花米草入侵我国海滩芦苇种群的机制之一。     

不同水分条件下混种密度对芦苇和反枝苋种间竞争的影响

与本地植物的种间竞争是影响外来植物能否成功入侵的关键因素之一,该研究通过受控模拟试验研究了本地植物芦苇(Phragmites australis)和外来入侵植物反枝苋(Amaranthus retroflexus)在淹水和干旱两种水分条件下混种密度(6∶2、4∶4和2∶6)对其种间竞争的影响。结果表明:(1)芦苇和反枝苋的相对产量与相对产量总和均小于1,即两种植物存在种间竞争。(2)种间竞争使芦苇和反枝苋的生长均受到了不同程度的抑制,表现在两者的株高和生物量均随着竞争者密度的增加而降低。(3)植株地上部分和地下部分的氮浓度表现出与株高和生物量相同的趋势,且在不同水分条件下存在差异。(4)芦苇和反枝苋分别在淹水和干旱环境下具有较强竞争力,但在各自较高混种密度下亦具有较强竞争力。可见,芦苇和反枝苋的种间竞争受到了水分和混种密度的影响。因此,在有反枝苋分布的湿地中,植物生长初期可通过增加土壤水分和/或增加芦苇等本地植物的种群密度以降低反枝苋的种群密度来限制其竞争能力,防止反枝苋在湿地中生长建群和扩散入侵。     

紫茎泽兰与黑麦草苗期相对竞争力和生长特征的研究

为探究替代物种黑麦草（Lolium perenne）抑制外来人侵杂草紫茎泽兰（Eupatorium adenophorum）的潜力,采用de Wit取代系列设计,在野外小区内设置3水平多年生黑麦草栽植时间、4水平种植密度以及3水平混种比例等不同种植处理,于12周后收获,分别测量黑麦草与紫茎泽兰的地上生物量和地下生物量,用单株相对产量分析其竞争力。结果表明,种植密度、混种比例和黑麦草栽植时间显著影响紫茎泽兰的地上生物量,紫茎泽兰的竞争力在低种植比例情况下,随黑麦草栽植时间的推迟而明显增强,随种植密度的降低而增强。种植密度、混种比例、黑麦草栽植时间以及这些因素的互作等对黑麦草的地上生物量与地下生物量均具有显著影响;栽植时间越晚、密度越大,黑麦草的竞争力就越弱（尽管不存在直线关系）。在大多数种植处理条件下,紫茎泽兰经历的种间竞争明显大于种内竞争,表现出较弱的竞争力;而黑麦草在与紫茎泽兰同时栽植的情况下,经历的种内竞争大于种间竞争,表现出较强的竞争力。     

氮肥和种植密度对紫茎泽兰生长和竞争的影响

紫茎泽兰（Ageratina adenophora）是我国西南热带亚热带地区危害最严重的外来入侵植物之一。本研究在野外设置不同氮肥供给水平和种植密度下的紫茎泽兰、拔毒散（Sida szechuensis）、伏生臂形草（Brachiaria decumbens）和非洲狗尾草（Setaria anceps）的单种和紫茎泽兰与其他3种植物混种的人工群落,从植被-土壤相互作用的角度分析紫茎泽兰的入侵机制。结果表明：单种时4种受试植物均表现了对氮营养的响应,随氮肥水平的升高,根冠比降低,比叶面积升高;通过比较总生物量增幅,氮响应能力非洲狗尾草>伏生臂形草>紫茎泽兰>拔毒散。混种时受试植物与紫茎泽兰的生物量比和株高也表明非洲狗尾草的竞争能力>伏生臂形草>紫茎泽兰>拔毒散;与拔毒散混种的紫茎泽兰以及与 紫茎泽兰混种的伏生臂形草和非洲狗尾草的种内竞争随种植密度的增大而增大,导致种间竞争能力下降,在高种植密度下对混种物种的抑制作用减弱。紫茎泽兰在氮响应能力、竞争能力和表型可塑性方面均强于本地种,可能与其入侵机制有关;牧草的氮响应能力和竞争能力强于紫茎泽兰,其中非洲狗尾草又强于伏生臂形草,可能更适宜于入侵地的替代控制。     

绿肥与粮,油作物间套混种的共生习性及再生机制的研究

绿肥与粮、油作物间套混种的共生习性及再生机制的研究包兴国,吕福海,桑同山,刘生战（甘肃省农科院土肥所,兰州７３００７０）Ｃｏ－ＧｒｏｗｔｈＨａｂｉｔａｎｄＲａｔｏｏｎｉｎｇＭｅｃｈａｎｉｓｍｏｆＧｒｅｅｎＭａｎｕｒｅＣｒｏｐｓＩｎｔｅｒ－ｏｒＭｉｘｅ...     

两种入侵植物与三种本地植物根系特征的比较研究

以菊科2种入侵植物飞机草(Chromolaena odorata(L.)R.M.King&H.Rob)和紫茎泽兰(Eupatorium adenophorum Speng)以及生活型相似的3种本地植物异叶泽兰(Eupatorium heterophyllum DC.)、佩兰(Eupatorium fortune Turcz.)和白头婆(Eupatorium japonicum Thunb.)为研究对象,以单种为对照,分析5种植物的株高、生物量以及细根形态和结构特征对混种(竞争)的响应。结果显示,在单种情况下,2种入侵植物的株高和地上生物量比3种本地植物高。2种入侵植物的株高和地上生物量在单种和混种下无显著差异,但在混种下3种本地植物的株高、地上生物量、根生物量、细根长度、细根表面积和体积与单种相比显著降低。在混种情况下,2种入侵植物的根组织密度显著降低,但比根长和比根面积仅在紫茎泽兰中显著增加。5种植物在单种和混种下的株高与细根长度、根生物量呈显著正相关,地上生物量与细根体积、表面积呈显著正相关。主成分分析结果表明入侵植物位于株高和地上生物量更高的一端。说明在与本地植物竞争的过程中,飞机草和紫茎泽兰能通过调整根系碳投资策略实现更强的竞争力。     

单种及混合种植对花叶芦竹铅积累量的影响

该研究采用盆栽试验法,选取石菖蒲、水生鸢尾、美人蕉、花叶芦竹为试验材料,将石菖蒲、水生鸢尾、美人蕉与花叶芦竹混种,用花叶芦竹单种作为对照,研究了混种对花叶芦竹生物量、根冠比以及铅积累量的影响。结果表明:(1)在不同浓度铅污染处理下,与花叶芦竹单种相比,混种模式增加了花叶芦竹的根冠比,且随着铅污染浓度的增加,表现出先上升后下降的趋势。(2)在单种和混种模式下花叶芦竹体内和土壤内的铅含量均表现出明显的浓度积聚效应;在混种模式下植物体内和土壤内的铅含量明显低于单种模式,且与单种模式之间的差异性显著(P0.05)。(3)在高浓度铅污染处理下,单种和混种模式的花叶芦竹铅转运系数均大于1,而富集系数则表现为在低浓度和高浓度铅污染处理下均大于1;在美人蕉+花叶芦竹的混种模式中,花叶芦竹的地上部分和地下部分的铅富集系数达到最大。因此,我们认为美人蕉+花叶芦竹混种模式,显著提高了花叶芦竹的生物量和根冠比,并对铅污染土壤的净化效果最佳。     

玉米产量对密度的敏感性研究

作物的密度敏感性反映了稳产性,是广义作物耐密性的重要组成部分.密度敏感性可以用敏感度表示,它是当密度偏离最适值1株/m^2时,产量-密度的单峰曲线上两条切线的到角,取值0°~180°,据此将密度敏感性划分为5个等级.密度敏感度的上界是最适单株产量的严格增函数.对中国上世纪60年代以来155个玉米密度试验产量结果作密度敏感性分析,发现密度敏感度接近对数正态分布,中位数为49°,极不敏感和不敏感两个等级占总数的68.4%.密度敏感度揭示了密度与稳产性的关系,在玉米及其它作物研究中具有重要应用价值.     

模拟人工湿地中植物多样性对铵态氮去除的影响

为了研究植物多样性对人工湿地生态系统的氮去除功能和硝化作用的影响,在模拟人工湿地中配置了单种和4种植物混种2个处理,并以铵态氮为唯一入水氮源负荷。结果表明:混种系统出水中的无机氮浓度显著低于单种系统(分别为3.41和7.20mg·L-1,P<0.05),铵态氮浓度也显著低于单种系统(分别为1.35和4.11mg·L-1,P<0.05);而出水硝铵浓度比(NO3-∶NH4+=1.55)则高于(P<0.05)单种系统(0.80),说明多样性增强了系统的硝化作用;混种系统基质的无机氮存留量(1455mg·m-2)低于(P<0.05)单种(2235mg·m-2),说明混种系统中可能存在资源的互补利用;根据物质平衡法推算出混种系统中植物总的氮吸收量对无机氮去除的贡献率(48%)大于(P<0.05)单种(31%),植物的可移除部分(地上)对无机氮移除的贡献率也呈现此规律(混种和单种分别为33%和20%,P<0.05);基质氮存留在2种系统中的贡献率则与植物吸收规律相反(混种和单种分别为5%和9%,P<0.05);混种系统中的反硝化作用、氨挥发和微生物的氮固持等对氮去除的贡献率低于单种系统。     

Effects of mixed cropping on population densities and parasitism rates of the diamondback moth, <Emphasis Type="Italic">Plutella xylostella</Emphasis> (Lepidoptera: Plutellidae)

Effects of mixed cropping and barrier crops on the population density and parasitism of the diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), were evaluated in field plots of cabbage grown in Bali, Indonesia. The densities of P. xylostella at larval and pupal stages, as well as the overall density at larval plus pupal stages, were significantly lower in cabbage/coriander mixed cropping subplots than in cabbage monoculture subplots. Parasitism of P. xylostella by the larval parasitoid Diadegma semiclausum (Hellen) (Hymenoptera: Ichneumonidae) was not significantly different between the mixed and monocultural cropping systems. These results do not support the so-called enemies hypothesis, but suggest that disruption of the host searching behavior of female moths by neighboring non-host plants is the mechanism behind the associational resistance observed in the coriander mixed cropping system. The inclusion of a Napier grass barrier between mixed crop and monoculture subplots did not affect the influence of mixed cropping on larval and pupal densities. Therefore, Napier grass, which is used locally as a fence for preventing livestock invasion of fields, would not obstruct the pest-reducing effect of coriander/cabbage mixed cropping.     

Modelling herbivore movement and colonization: pest management potential of intercropping and trap cropping

1 Using a stochastic simulation model, we explored the effects of agroecosystem diversity on herbivore densities. 2 Using parameters that included reproduction, colonization, and local movement rates, we simulated an insect herbivore population colonizing rows of plants in an agricultural setting. 3 Plant rows were comprised of either principal crop, intercrop, or trap crop. Herbivore parameters varied for different plant types. 4 Percent crop cover and movement rates were varied, and ensuing herbivore densities on crop rows were recorded. 5 In trap cropping schemes, both percent crop cover and movement rates were critical in determining herbivore densities. Intercropping schemes were governed primarily by colonization rates. 6 These results suggest that trap cropping schemes merit more attention than intercropping systems in the design and analysis of mixed cropping systems.     

Population Dynamics of the Root-Knot Nematode Meloidogyne Triticoryzae under Five Rice-Based Cropping Systems

Population dynamics of M. triticoryzae in five rice-based cropping systems have revealed that rice was the main host. Most of the population growth which contributed to the peak population density in September occurred in the rice crop. The other crops in the Rabi and summer seasons had a profound influence on the height of the peak in the succeeding rice crop. Some level of reproduction occurred in wheat, greengram and berseem in Rabi which contributed to relatively higher equilibrium densities. The potato variety Pusa Bahar increased the population density while Pusa Badshah decreased it. The lower temperature during the Rabi season also limited the rate of population growth. The incorporation of organic matter into the soil, either as crop residues of greengram or berseem, reduced the population growth of M. triticoryzae .     

Agricultural land use and Skylark Alauda arvensis: a case study linking a habitat association model to spatially explicit change scenarios

The development of forward scenarios is a useful method of envisaging the environmental implications of potential changes in land use, as a tool for policy development. In this paper, a spatially explicit case study is used to provide insight into the environmental impacts of Common Agricultural Policy reform on Skylark Alauda arvensis, a species which is widespread on arable farmland, breeds in crops and has declined in recent decades. A generalized linear mixed model was used to estimate Skylark breeding population densities in different crops, using survey data collected from farms in the east of England, supplemented by the literature. Model outputs were then used to predict Skylark densities in an East Anglian Joint Character Area dominated by arable cropping. Predicted densities were mapped at field level using GIS, based on actual cropping derived from Integrated Administration and Control System data collected for the administration of subsidy payments. Three future scenarios were then created, based on expert opinion of potential changes in cropping over the next 5 years, and potential changes in Skylark density mapped on the basis of the predicted changes in cropping patterns. Overall, Skylark densities were predicted to decrease on average by 11–14% under ‘market‐led’ (increasing wheat and oilseed rape, reduced set‐aside) and ‘energy crop’ (5% area under short rotation coppice) scenarios, but remained virtually unchanged under an ‘environment‐led’ (diverse cropping) scenario. The ‘market‐led’ scenario is closest to short‐term agricultural trajectories, but wider cultivation of biomass energy crops as modelled under the ‘energy crop’ scenario could occur in the medium term if energy policies are favourable. Appropriate mitigation strategies therefore need to be implemented if a continued decline in the Skylark population on lowland arable farmland is to be averted. The results provide a readily accessible visualization of the potential impacts of land‐use change for policy‐makers; similar techniques could be applied to visualize effects of changes arising through other drivers, including climate change.     

Strategies to decrease nitrate leaching in the Brimstone Farm Experiment,Oxfordshire, UK, 1988–1993: the effects of winter cover crops and unfertilised grass leys

Nitrate losses in drainflow were measured over five years on eight hydrologically isolated field plots, pairs of which had the following cropping regimes: (a) a 3-yr unfertilised, ungrazed grass ley followed by winter and spring cereals, (b) mixed cropping including winter cover crops, spring cereals, winter cereals, winter fallow and spring beans, (c) a similar sequence to (b) but with a winter fallow replacing the cover crop in the first year and a winter cover crop replacing the fallow in the third year, and (d) continuous winter cereals (control plots). Less nitrate was lost in winter drainflow from winter cover crops than from the winter fallows, but over all five years less nitrate was leached from the continuous cereal plots than from those with mixed cropping. Most of the extra nitrate lost from the mixed cropping regimes probably resulted from mineralisation of the cover crop residues, which occurred at times when subsequent crops could not take advantage of the mineral nitrogen released. Crops grown after the grass ley and cover crops did not benefit from their residues, in terms of either grain yield or of total nitrogen uptake. We conclude that on heavy clay soils in UK a cropping regime of continuous winter cereals offers the best compromise between profitable crop production and minimised nitrate loss to surface waters.     

Crop identity and memory effects on aboveground arthropods in a long‐term crop rotation experiment

Agricultural landscapes are globally dominated by monocultures under intensive management. This is one of the main reasons for biodiversity loss and insect population decline in many regions all over the world. Agroecosystem biodiversity in these areas can be enhanced by cropping system diversification, such as crop rotations. Yet, long‐term studies on effects of crop rotations on aboveground agrobiodiversity are lacking. We set up a 10‐year long‐term crop rotation experiment in Central Germany and monitored the temporal dynamics of aboveground arthropods over a full cultivation period to investigate influence of current and preceding crop identity and cropping system diversification on activity density, species richness, and community structure. We found that species composition was strongly influenced by currently grown crop although effect on arthropods varied between species groups. Especially, winter oilseed rape strongly affects arthropod community structure. Interestingly, we were also able to show an influence of the preceding crops, indicating an ecological memory effect in the aboveground arthropod community. Our results show that crop identity of both currently and previously grown crops in crop rotations may lead to an increase in arthropod activity density and changes in species composition. Diversified crop rotations including appropriate crops can be an easily implemented tool to increase arthropod biodiversity and biomass at large spatial and temporal scales, particularly in areas dominated by a single crop (e.g., wheat, maize). Our results may help to design optimized crop rotations for large‐scale enhancement of insect biodiversity in agroecosystems.     

Agricultural landscapes with organic crops support higher pollinator diversity

Pollinators are traditionally thought to perceive non-flowering crop fields as hostile landscape matrix. In this study, we show that landscapes composed of higher proportions of organic crop fields support more bee species at greater abundances in fallow strips. An increase in organic cropping in the surrounding landscape from 5% to 20% enhanced bee species richness in fallow strips by 50%, density of solitary bees by 60% and bumble bee density by 150%. Bee species richness and bumble bee density responded strongest to organic cropping in landscape sectors with 500 m radius, solitary bee density in landscape sectors with 250 m radius. The most likely source of these results is that crop and noncrop habitats are strongly connected via bee foraging at the landscape scale. It seems likely that bees depending on nesting sites in fallow strips benefited from the more abundant flower resources provided by broadleaved weeds in organic crop fields. We conclude that the incorporation of organic crop fields into conventionally managed agricultural landscapes can provide food resources needed to sustain greater pollinator species richness in noncrop habitats.     

Strip-tilled Cover Cropping for Managing Nematodes, Soil Mesoarthropods, and Weeds in a Bitter Melon Agroecosystem

A field trial was conducted to examine whether strip-tilled cover cropping followed by living mulch practice could suppress root-knot nematode (Meloidogyne incognita) and enhance beneficial nematodes and other soil mesofauna, while suppressing weeds throughout two vegetable cropping seasons. Sunn hemp (SH), Crotalaria juncea, and French marigold (MG), Tagetes patula, were grown for three months, strip-tilled, and bitter melon (Momordica charantia) seedlings were transplanted into the tilled strips; the experiment was conducted twice (Season I and II). Strip-tilled cover cropping with SH prolonged M. incognita suppression in Season I but not in Season II where suppression was counteracted with enhanced crop growth. Sunn hemp also consistently enhanced bacterivorous and fungivorous nematode population densities prior to cash crop planting, prolonged enhancement of the Enrichment Index towards the end of both cash crop cycles, and increased numbers of soil mesoarthropods. Strip-tilled cover cropping of SH followed by clipping of the living mulch as surface mulch also reduced broadleaf weed populations up to 3 to 4 weeks after cash crop planting. However, SH failed to reduce soil disturbance as indicated by the Structure Index. Marigold suppressed M. incognita efficiently when planted immediately following a M. incognita-susceptible crop, but did not enhance beneficial soil mesofauna including free-living nematodes and soil mesoarthropods. Strip-tilled cover cropping of MG reduced broadleaf weed populations prior to cash crop planting in Season II, but this weed suppression did not last beyond the initial cash crop cycle.     

苹果园植被多样化在果树害虫持续治理中的作用

1992-1995年对四个不同类型苹果园进行了系统调查,结果表明引入夏至草、泥胡菜等有益杂草并与树行间种植的紫花苜蓿搭配的生草园,捕食性天敌发生量大,种群稳定,发生高峰期较免耕园、清耕园提前7d左右;害虫发生量低,种群数量变化平稳,无突增现象,生草园在未施用任何杀虫、杀螨剂的情况下,树冠上蚜虫、叶螨和潜叶蛾的平均虫口密度均低于免耕园、清耕稀植园和密植园。生草园寄生性天敌的数量亦明显增加,金纹细蛾的寄生率高达62.7%,分别是免耕园、清耕稀植和密植园的1.34倍、16.9倍和17.9倍。果园植被多样化可使地面植被上的天敌数量逐年成倍提高,生草第3年捕食性天敌的数量是生草当年的12.5倍,树冠上天敌量亦明显增加,益害比大大提高,生草第2年树冠上最大蝽螨比为1∶14.4,与原先油菜与紫花苜蓿搭配生草5年的效果相近。除当年生草果园(1992年)早期施用1次杀螨剂—螨克外,1993、1994年均未施用任何杀虫、杀螨剂,通过天敌的自然控制作用,有效地控制果树害螨、蚜虫和金纹细蛾等次生性害虫的为害。     

不同种植方式下转Bt基因水稻对稻田节肢动物群落的影响

如何设置庇护所使其在转基因作物靶标害虫抗性治理中达到更高的效率,就此,本文 研究了Bt水稻与非Bt水稻不同种植方式对Bt稻田节肢动物群落的影响。结果表明,整体上,块块种植和列列混栽方式对Bt稻田节肢动物群落多样性、均匀度和优势集中性无显著影响;但混栽方式下的Bt稻田节肢动物群落个体密度和物种丰富度显著偏低。在水稻发育中期,混栽方式下Bt稻田节肢动物群落季节动态与块块种植方式下的明显不同,其多样性、均匀度、优势集中性指数均类似于同种植方式下非Bt稻上各参数的季节动态。混栽方式下的庇护所效应更明显。