镁碱化盐土微生物生物量和土壤基础呼吸

通过测定甘肃河西走廊疏勒河中游昌马冲积扇缘不同镁碱度条件下10个采样点30个土样的化学性质和生物化学性质指标,研究了电导率和镁碱度对土壤微生物生物量及其基础呼吸的影响。结果表明:微生物生物量碳(氮)和土壤基础呼吸与电导率、镁碱度和Mg2+/Ca2+之间显著负相关,表明盐度和镁碱度对土壤微生物群落有显著的抑制作用,而且盐度的抑制作用比镁碱度更大;微生物代谢熵(qCO2)和电导率、镁碱度、Mg2+/Ca2+之间为正相关关系,也说明镁碱化盐土对土壤微生物而言是一种严重的胁迫环境。     

长期施肥处理对玉米生长期潮土微生物生物量和活度的影响

以1989年建立的中国科学院封丘农田生态系统国家试验站的长期定位试验为平台,研究经18a连续不同施肥处理后玉米季土壤微生物生物量碳氮和微生物活度的动态变化及其与土壤有机碳之间的相互关系,并探讨施肥措施对土壤微生物及其活性的影响。施肥处理包括:(1)有机肥(OM);(2)1/2化肥和1/2有机肥(1/2OM+1/2NPK);(3)氮磷钾肥(NPK);(4)氮磷肥(NP);(5)磷钾肥(PK);(6)氮钾肥(NK);(7)不施肥,即对照(CK)7个处理。结果表明,微生物生物量碳氮和微生物活度在玉米生长期内均有明显的时间变异性,其中微生物生物量碳与微生物活度的动态变化比较一致,其间的极显著相关关系表明潮土微生物生物量碳的变化可以在很大程度上代表土壤微生物活度的变化。施肥制度显著影响微生物生物量碳氮和微生物活度的变化,总体趋势为OM1/2OM+1/2NPKNPKNPPKNKCK,表明OM有利于保持土壤的生物化学环境及促进土壤的生物学活性;与OM处理相比,化学肥料的长期施用有降低土壤微生物生物量和微生物活度的趋势,尤其是缺素处理的表现更为明显,其中以缺磷处理的表现最为严重。土壤微生物生物量碳氮、微生物活度与土壤有机碳变化均呈极显著正相关。     

三峡库区紫色土旱坡地桑树配置模式对土壤微生物生物量碳氮的影响

为探讨评价三峡库区紫色土旱坡地土壤肥力及土壤质量的生物学指标,本试验研究了三峡库区典型水土保育模式——桑树护坡对土壤微生物生物量碳氮的影响。试验共设横坡农作、双边桑树+横坡农作、等高桑树+双边桑树+横坡农作及四边桑树+等高桑树+横坡农作4个处理。结果表明,不同坡长及桑树布局间土壤微生物生物量碳氮差异显著;双边桑树+横坡农作处理能够显著降低土壤微生物生物量氮对土壤氮库的贡献,使土壤微生物生物量碳平均提高23.43 mg·kg-1;而四边桑树+等高桑树+横坡农作处理却能够显著降低土壤微生物商,使土壤微生物生物量氮平均提高21.81 mg·kg-1。此外,在横坡农作处理中土壤微生物生物量碳氮与土壤碳氮具有极显著相关性,而在旱坡地桑树系统中,土壤微生物生物量碳氮与土壤碳氮相关性大多不显著。由此可知,旱坡地农桑配置模式在一定程度上可以提高土壤微生物生物量碳氮,但不同桑树布局对土壤微生物生物量碳氮在三峡库区紫色土旱坡地桑树系统中的维持能力并不一致,将微生物生物量碳氮作为紫色土旱坡地土壤质量演变的评价指标值得商榷。     

阿特拉津除草剂对土壤微生物生态特征的影响

采用实验室培养试验研究了阿特拉津除草剂对土壤微生物生态特征的影响。结果表明：阿特拉津除草剂10mg／kg浓度处理释放的CO2与对照无明显差异,根据危害系数法的分级方法,其属于无实际危害级农药;阿特拉津除草剂10mg／kg浓度处理在初始阶段,明显降低土壤微生物生物量碳和微生物生物量氮,随培养时间推移土壤微生物生物量碳和生物量氮有所恢复,随后又有所下降,30d后土壤微生物生物量变化趋于平缓;阿特拉津除草剂对土壤微生物多样性产生一定影响．其影响随培养时间而异。在一定程度上反映出土壤微生物生态特征指标可作为除草剂污染土壤环境质量变异的参考指标。     

生物质炭长期施用对旱地红壤微生物特性及理化性质的影响

为了解生物质炭施于旱地红壤较长时间后对土壤物理、化学及微生物特性的影响,以江西典型旱地红壤为对象,采用田间长期定位试验的方法观察了不同用量生物质炭(0、2.5、5、10、20、30 t hm~(-2)和40 t hm~(-2))在施用4年后旱地红壤基础理化性质、土壤微生物生物量碳、氮、土壤基础呼吸强度及微生物商、微生物代谢熵等的变化。结果表明,施用生物质炭4年后,旱地红壤pH、有机碳、全氮、微生物生物量碳和生物量氮均随生物质炭施用量的增加呈上升趋势,土壤容重、微生物碳氮比呈下降趋势。以生物质炭用量40 t hm~(-2)时的改良效果最好,与对照相比,土壤容重显著(P0.05)降低了0.17 g cm-3,微生物碳氮比显著降低了7.97,土壤pH、有机碳、全氮、微生物生物量碳和生物量氮分别显著提高了6.1%、47%、21.5%、43.3%和162.6%。试验证明施用生物质炭较长时间后对旱地红壤的改良效果依旧良好,并且生物质炭施用量越高,对土壤理化性质和微生物特性的影响越显著。     

长期免耕与施用有机肥对土壤微生物生物量碳、氮、磷的影响

通过设置在江苏省句容农科所的田间定位试验研究长期免耕及施用有机肥料对土壤微生物生物量碳、氮、磷的影响。结果表明 :经过 1 6年 32茬稻—麦水旱轮作后 ,表土层 ( 0～ 5cm)土壤微生物生物量碳、氮、磷含量比亚表层 ( 5～ 1 0cm)分别高 2 7.5 %、43.6%和1 1 %。与常规耕翻相比长期免耕处理表土层土壤微生物生物量碳、氮含量分别增加了2 5 .4%和 45 .4% ,而微生物生物量磷无明显变化规律 ;亚表层的土壤微生物生物量碳、氮、磷免耕与耕翻两种耕作方式间的差异不显著。尽管各施肥处理施用的氮、磷、钾数量完全相等 ,但土壤微生物生物量碳、氮、磷的含量却因肥料种类的不同而异。综合 0～ 5和 5～ 1 0cm土层 ,微生物生物量碳、磷为 :猪粪 化肥 >秸秆 化肥 >绿肥 化肥 >化肥 >不施肥 ,微生物生物量氮则为 :猪粪 化肥 >绿肥 化肥 >秸秆 化肥 >化肥 >不施肥。相关分析结果显示 ,土壤微生物生物量碳、氮与土壤有机碳、土壤全氮和土壤碱解氮之间均呈极显著的正相关 ,表明其与土壤肥力关系密切 ,可作为评价土壤肥力性状的生物学指标     

添加硝化抑制剂DMPP对红壤水稻土硝化作用及微生物群落功能多样性的影响

通过室内培育试验,研究了不同施氮水平下添加硝化抑制剂(DMPP)处理对红壤水稻土NH4+-N、NO3–-N含量、微生物生物量碳及微生物群落功能多样性的影响。结果表明:56天培养期内,不同处理的NH4+-N含量总体呈下降趋势,而NO3–-N含量呈上升趋势。随施氮水平提高,培养期内NH4+-N平均含量从0 mg/kg处理的24.10 mg/kg增加到400 mg/kg处理的412.10 mg/kg,NO3–-N平均含量从0 mg/kg处理的41.88 mg/kg增加到400 mg/kg处理的99.83 mg/kg。添加DMPP显著抑制硝化作用进行,抑制效果随施氮量增加而提高,400 mg/kg施氮水平下,添加DMPP硝化率和硝化速率比不添加DMPP处理分别下降了29.0%和44.3%,下降幅度远大于其他施氮水平处理。施氮水平也影响土壤微生物生物量碳和微生物群落功能多样性。施氮量从0 mg/kg增加到400 mg/kg,土壤微生物生物量碳下降了12.5%,AWCD值下降了78.4%,Shannon指数下降了22.3%;与不添加DMPP处理相比,添加DMPP处理的土壤微生物生物量、AWCD值、Shannon指数分别提高了2.1%、23.9%、7.8%,尤其在400 mg/kg施氮水平下,提高的幅度更加明显。     

不同施肥水平及玉米种植对土壤微生物生物量碳氮的影响

以裸地(不种植作物)和玉米种植的田间小区试验为平台,研究不同施肥水平及玉米种植对土壤微生物生物量碳氮的影响。结果表明,当不施肥时,土壤微生物生物量碳氮含量裸地平均值分别为175.98 mg/kg和26.04 mg/kg,种植玉米小区的平均值分别为161.65 mg/kg和22.70 mg/kg,土壤微生物生物量碳氮低于裸地;而施肥时,裸地的土壤微生物生物量碳氮平均值的变化范围分别为182.27~206.27 mg/kg和27.41~31.22 mg/kg,种植玉米小区的变化范围分别为194.70~235.58 mg/kg和35.76~44.66 mg/kg,土壤微生物生物量碳氮高于裸地,可见土壤碳氮的平衡对于土壤微生物生物量碳氮极为重要。裸地和玉米种植小区土壤微生物生物量碳氮均随着施肥量的增加呈现出先增加后降低的趋势,其施氮水平拐点分别为70 kg/hm2和150kg/hm2,表明施肥水平对土壤微生物生物量碳氮具有显著影响。另外,玉米各生育期间土壤微生物生物量碳氮也存在着显著差异,其中,土壤微生物生物量碳氮含量在拔节期处于最低,变动范围分别为154.46~229.09 mg/kg和18.84~31.44 mg/kg;抽雄期处于最高,变动范围分别为171.71~242.48 mg/kg和30.01~50.54 mg/kg。     

添加秸秆碳源对土壤微生物生物量和原生动物丰富度的影响

为了研究引入秸秆碳源对根结线虫(Meloidogyne spp.)病害严重土壤中微生物生物量和原生动物的影响, 以番茄为供试作物, 设置4个梯度的小麦秸秆添加量[CK(0 g·kg^-1), 1N(2.08 g·kg^-1)、2N(4.16 g·kg^-1)和4N(8.32 g·kg^-1)], 研究不同种植时间(6个月和4个月)下土壤微生物生物量碳、氮和原生动物丰度的变化。研究结果表明: 添加秸秆对微生物生物量碳、氮和原生动物丰富度有明显促进作用, 添加的秸秆量越多, 这种促进作用越明显。不同秸秆添加量处理中, 微生物生物量碳、氮和原生动物丰度为: 4N>2N>1N>CK。秸秆对原生动物的群落结构也有显著影响, 在各处理中, 鞭毛虫和肉足虫占有绝大比例, 分别占总丰度的29.44%和66.19%, 纤毛虫仅占4.37%。在相同添加秸秆量条件下, 土壤原生动物丰度随种植时间的延长而提高, 而微生物生物量碳、氮量随种植时间的延长而降低。而在种植时间相同条件下, 随着秸秆量的增加土壤微生物生物量碳、氮量和微生物生物量碳氮比和原生动物总丰度相应增加。     

存在有机酸时pH对镉毒害土壤微生物生物量的影响

采用室内培养实验方法,研究了存在有机酸时ＰＨ对镉毒害土壤微生物生物量的影响。结果表明,存在一定量有机酸的镉污染土壤中,土壤中微生物生物量碳（Ｃｍｉｃ）和氮（Ｎｍｉｃ）随着ＰＨ升高而升高,其中含胡和线条到为土壤中微生物生物量碳（Ｃｍｉｃ）和氮（Ｎｍｉｃ）明显比含低分子量有机酸（柠檬要到、酒石酸、乙酸、草酸）的土壤增加的快,土壤中微生物生物量碳氮比（Ｃｍｉｃ／Ｎｍｉｃ）随或高而降低,表明提高土壤ＰＨ可     

Microbial biomass and activity in alkalized magnesic soils under arid conditions

The effects of salinity and Mg²⁺ alkalinity on the size and activity of the soil microbial communities were investigated. The study was conducted along the border area of the alluvial fan of the Taolai River. Thirty soil samples were taken which had an electrical conductivity (EC) gradient of 0.93-29.60 mS cm⁻¹. Soil pH ranged from 8.60 to 9.33 and correlated positively with Mg²⁺/Ca²⁺ ratio, exchangeable Mg²⁺ percentage and HCO₃⁻+CO₃²⁻. Mg²⁺/Ca²⁺ varied considerably from 3.04 to 61.31, with an average of 23.03. Exchangeable Mg²⁺ percentage generally exceeded 60% and had a positive correlation with Mg²⁺/Ca²⁺. HCO₃⁻+CO₃²⁻ averaged 1.63 cmol kg⁻¹ and usually did not exceed 2.0 cmol kg⁻¹.Microbial biomass, indices of microbial activity and the activities of the hydrolases negatively correlated with Mg²⁺/Ca²⁺ or exchangeable Mg²⁺ percentage. Biomass C, biomass N, microbial quotient (the percentage of soil organic C present as biomass C), biomass N as a percentage of total N, potentially mineralizable N, FDA hydrolysis rate and arginine ammonification rate decreased exponentially with increasing EC. The biomass C/N tended to be lower in soils with higher salinity and Mg²⁺ alkalinity, probably reflecting the bacterial dominance in microbial biomass in alkalized magnesic soils. The metabolic quotient (qCO₂) positively correlated with salinity and Mg²⁺ alkalinity, and showed a quadratic relationship with EC, indicating that increasing salinity and Mg²⁺ alkalinity resulted in a progressively smaller, more stressed microbial communities which was less metabolically efficient. Consequently, our data suggest that salinity and Mg²⁺ alkalinity are stressful environments for soil microorganisms.     

Active organic matter distribution in the surface 15 cm of undisturbed and cultivated soil

Summary Organic-matter dynamics near the soil surface influence plant nutrient supplies. To evaluate the effects of cultivation on active soil organic-matter distribution in the surface 15 cm, we measured total organic C, Kjeldahl N, microbial biomass C and N (chloroform fumigation), respirable C and mineralizable N (aerobic incubations) in 1-cm layers from 0–10 cm and in 2.5-cm layers from 10–15 cm in adjacent areas of undisturbed (shortgrass steppe) and cultivated (four years wheat-fallow rotation) Ascalon sandy loam soil (Aridic Argiustoll). The active organic matter was highly concentrated in the surface centimeter of undisturbed soil. In undisturbed soil, microbial biomass C and N concentrations were more than five times greater at 0–1 cm than at 2–15 cm. Respirable-C and mineralizable-N concentrations (20-day incubations) were 8 and 18 times greater at 0–1 cm than at 2–15 cm. Below 3 cm, the concentrations were equal in both cultivated and undisturbed soil. Cultivation reduced the average concentrations at 0-15 cm of microbial biomass C and N by 62% and 32%, and of respirable C and mineralizable N by 71% and 46%, by reducing the concentrations at 0-1 cm to levels comparable to the 2- to 15-cm layers of the undisturbed soil. Decreases after cultivation ensued from disrupting the surface 1-cm layer.     

Soil organic matter quality as influenced by tillage,lime, and phosphorus

In Eastern Canada, cereal yields are often restricted by soil acidity and low fertility. Continuous cereal production can also lead to soil structural degradation. The addition of lime and fertilizers and the adoption of conversation tillage practices are proposed solutions which may have a positive impact on soil quality. The objective of the present work was to assess the impact of 3 years of different tillage practices and P additions, and of a single lime addition on organic C and total N, microbial biomass C, and on N mineralization at the surface layer (0–7.5 cm) of a Courval sandy clay loam (Humic Gleysol). The easily mineralizable N, total amount of N mineralized in 22.1 weeks, the rate of N mineralization, and microbial biomass C were significantly greater in the minimum tillage than in the moldboard plow treatment. Chisel plow treatment showed intermediate values. The ratios of potentially mineralizable N and of easily mineralizable to total soil N were also significantly larger under minimum tillage and chisel plowing than under moldboard plowing. The lime and P treatments had no significant effect on the measured soil quality parameters. The total amount of N mineralized per unit of biomass C decreased as the tillage intensity increased, suggesting a decrease in the efficiency of the biomass in transforming organic N into potentially plant-available forms and thus a loss in soil organic matter quality. The results of this study indicate that conservation tillage practices such as rototilling and chisel plowing are efficient ways of maintaining soil organic matter quality when old pastures are brought back into cultivation.     

Soil microbial biomass and activity in organic tomato farming systems: Effects of organic inputs and straw mulching

Organic farming is rapidly expanding worldwide. Plant growth in organic systems greatly depends on the functions performed by soil microbes, particularly in nutrient supply. However, the linkages between soil microbes and nutrient availability in organically managed soils are not well understood. We conducted a long-term field experiment to examine microbial biomass and activity, and nutrient availability under four management regimes with different organic inputs. The experiment was initiated in 1997 by employing different practices of organic farming in a coastal sandy soil in Clinton, NC, USA. Organic practices were designed by applying organic substrates with different C and N availability, either in the presence or absence of wheat-straw mulch. The organic substrates used included composted cotton gin trash (CGT), animal manure (AM) and rye/vetch green manure (RV). A commercial synthetic fertilizer (SF) was used as a conventional control. Results obtained in both 2001 and 2002 showed that microbial biomass and microbial activity were generally higher in organically than conventionally managed soils with CGT being most effective. The CGT additions increased soil microbial biomass C and activity by 103-151% and 88-170% over a period of two years, respectively, leading to a 182-285% increase in potentially mineralizable N, compared to the SF control. Straw mulching further enhanced microbial biomass, activity, and potential N availability by 42, 64, and 30%, respectively, relative to non-mulched soils, likely via improving C and water availability for soil microbes. The findings that microbial properties and N availability for plants differed under different organic input regimes suggest the need for effective residue managements in organic tomato farming systems.     

Extractability of labelled microbial biomass N by electro-ultrafiltration and CaCl2 extraction

A laboratory soil incubation and a pot experiment with ryegrass were carried out in order to examine the extractability of microbial biomass N by using either 10-mM CaCl₂ extraction or the electro-ultrafiltration (EUF) method. The aim of the experiment was to test the hypothesis whether the organic N (Norg) extracted by EUF or CaCl₂ from dried soil samples represents a part of the microbial biomass. For the laboratory incubation a ¹⁵N-labelled Escherichia coli suspension was mixed with the soil. For the pot experiment a suspension of ¹⁵N-labelled bacteria was applied which had previously been isolated from the soil used. Soil samples of both treatments, with and without applied bacterial suspension, were extracted by EUF and CaCl₂. The extractability of applied microbial biomass was estimated from the difference in extractable Norg between the two treatments. In addition, the N isotopic composition in the upper plant matter, in the soil, and in organic and inorganic N fractions of EUF and CaCl₂ extracts was analysed. Both experiments showed that the applied microbial biomass was highly accessible to mineralization and thus represented potentially mineralizable N. However, this mineralizable N was not extractable by CaCl₂ or by the EUF method. It was, therefore, concluded that the organic N released on soil drying and which was thus extractable was derived from the non-biomass soil organic matter. The result suggests that both extraction methods may provide a suitable index for mineralizable N only in cases where the decomposable organic substrates are derived mainly from sources other than the living soil biota.Dedicated to Professor J. C. G. Ottow on the occasion of his 60th birthday     

Nitrogen pools and turnover in arable soils under different durations of organic farming: I: Pool sizes of total soil nitrogen,microbial biomass nitrogen,and potentially mineralizable nitrogen

Soil organic matter contents, soil microbial biomass, potentially mineralizable nitrogen (N) and soil pH values were investigated in the Ap horizons of 14 field plots at 3 sites which had been under organic farming over various periods. The objective was to test how these soil properties change with the duration of organic farming. Site effects were significant for pH values, microbial biomass C and N, and for potentially mineralizable N at 0—10 cm depth. The contents of total organic C, total soil N, and potentially mineralizable N tended to be higher in soils after 41 versus 3 years of organic farming, but the differences were not significant. Microbial biomass C and N contents were higher after 41 years than after 3 years of organic farming at 0—10 cm depth, and the pH values were increased at 10—27 cm depth. Nine years of organic farming were insufficient to affect soil microbial biomass significantly. Increased biomass N contents help improve N storage by soil micro‐organisms in soils under long‐term organic farming.     

Burning causes long-term changes in soil organic matter content of a South African grassland

The effects of burning a native grassland on soil organic matter status was investigated on a long-term (50 years) field experiment where different times and frequencies of burning were compared. Significant decreases in organic C were observed only in the surface 0-2 cm layer and only under annual and biennial winter burning and biennial and triennial autumn burning. Burning in spring did not significantly affect organic C content presumably because substantial amounts of litter decomposed and/or were incorporated into the soil by faunal activity prior to burning. Total N content was decreased substantially to a depth of 6 cm by all burning treatments and as a result, the C:N ratio of soil organic matter was widened. In addition, the amount of potentially mineralizable N, as measured by either aerobic incubation or plant N uptake in a pot experiment, was much reduced. Burning also induced a decrease in light fraction and hot water-extractable C in the 0-2 cm layer but an increase in these parameters, and in microbial biomass C and root density, in the 4-10 cm layer. This was attributed to burning causing a decrease in above-ground litter inputs but increased turnover of root material below the surface. Despite the decrease in organic C and total N content with increasing soil depth, potentially mineralizable N showed the opposite trend. This unexpected finding was confirmed at a nearby site under native grassland and contrasted with decreasing potentially mineralizable N with depth which was measured under a fertilized kikuyu grass dairy pasture. The wide C:N ratio of litter from native grassland, in association with the decreasing size and activity of the microbial biomass with depth results in greater N immobilization (thus less net mineralization) occurring in soil samples taken from close to the soil surface.     

Microbial biomass and mineralizable nitrogen distributions in no-tillage and plowed soils

Summary Distribution of soil microbial biomass and potentially mineralizable nitrogen (PMN) in long-term tillage comparisons at seven sites in the United States varied with tillage management and depth in soil. Microbial biomass and PMN levels of no-tillage soils averaged 54% and 37% higher, respectively, than those in the surface layer of plowed soil. Biomass and PMN levels were greatest in the surface 0 to 7.5-cm layer of no-tillage soil and decreased with depth in soil to 30 cm. Biomass and PMN levels of plowed soil, however, were generally greatest at the 7.5 –15 cm depth. Microbial biomass levels were closely associated with soil distributions of total C and N, water content, and water-soluble C as influenced by tillage management. Potentially mineralizable N levels in soil were primarily associated with distributions of microbial biomass and total N. Absolute levels of PMN and microbial biomass and the relative differences with tillage management were dependent on climatic, cropping, and soil conditions across locations. The additional N contained in soil biomass and PMN in the surface 0–7.5 cm of no-tillage compared with plowed soils ranged from 13 to 45 and 12 to 122 kg N/ha, respectively, for 6 of 7 locations. Fertilizer placement below the biologically rich surface soil layer and/or rotational tillage may improve short-term nitrogen use efficiency and crop growth on reduced-tillage soils.Contribution from USDA-ARS in cooperation with the Nebraska Agricultural Experimental Station, published as paper no. 8086, Journal Series, Nebraska Agricultural Experimental Station     

Earthworms increase soil microbial biomass carrying capacity and nitrogen retention in northern hardwood forests

Earthworms have been shown to produce contrasting effects on soil carbon (C) and nitrogen (N) pools and dynamics. We measured soil C and N pools and processes and traced the flow of ¹³C and ¹⁵N from sugar maple (Acer saccharum Marsh.) litter into soil microbial biomass and respirable C and mineralizable and inorganic N pools in mature northern hardwood forest plots with variable earthworm communities. Previous studies have shown that plots dominated by either Lumbricus rubellus or Lumbricus terrestris have markedly lower total soil C than uncolonized plots. Here we show that total soil N pools in earthworm colonized plots were reduced much less than C, but significantly so in plots dominated by contain L. rubellus. Pools of microbial biomass C and N were higher in earthworm-colonized (especially those dominated by L. rubellus) plots and more ¹³C and ¹⁵N were recovered in microbial biomass and less was recovered in mineralizable and inorganic N pools in these plots. These plots also had lower rates of potential net N mineralization and nitrification than uncolonized reference plots. These results suggest that earthworm stimulation of microbial biomass and activity underlie depletion of soil C and retention and maintenance of soil N pools, at least in northern hardwood forests. Earthworms increase the carrying capacity of soil for microbial biomass and facilitate the flow of N from litter into stable soil organic matter. However, declines in soil C and C:N ratio may increase the potential for hydrologic and gaseous losses in earthworm-colonized sites under changing environmental conditions.