Effects of elevated atmospheric CO2 and N fertilization on abundance,diversity and C-isotopic signature of collembolan communities in arable soil

Rising atmospheric CO₂ concentrations are expected to have marked impacts on the carbon (C) turnover in agro-ecosystems through increased plant photosynthetic rates, leading to an enhanced biomass, and wider plant C/N ratios. Through increased carbon allocation below-ground, as well as through changed litter quality, CO₂ enrichment will indirectly affect soil faunal communities. In the present study we investigated how elevated atmospheric CO₂ and two different levels of N fertilization may affect abundance and diversity of collembolans, as important catalysts in decomposition processes, within an agro-ecosystem under winter wheat cultivation. The investigations were carried out in 2002 within a field experiment using the “Free Air CO₂ Enrichment” technique (FACE) at the Federal Agricultural Research Centre (Braunschweig, Germany). Stable C-isotopic analysis of collembolans, soil, and crops gave insight into C translocation. During our investigations δ¹³C values of all components analysed were significantly more negative under FACE compared to ambient air conditions. Stable C-isotopic signatures of collembolans were similar to those of soil under ambient air, but in between those of soil and roots under elevated CO₂ conditions. Our results revealed significant effects of both treatments (CO₂ enrichment and N fertilization) on density and species diversity of collembolans. Overall, collembolans were stimulated under elevated CO₂ conditions, showing an increased abundance of more than 50% (11 240 ind m⁻²) as well as a higher biodiversity (Shannon Weaver index = 2.5; evenness = 0.75) compared to ambient air conditions (7520 ind m⁻²; Shannon Weaver index = 2.2; evenness = 0.72). With regard to N supply, a decrease of about 20–30% under CO₂ enrichment and 45–55% under ambient air conditions in collembolan abundance with no alteration in diversity was recorded under reduced N fertilization. The observed impacts were species-specific.     

Plant removal disturbance and replant mitigation effects on the abundance and diversity of low-arctic soil biota

Due to the dependence of soil organisms on plant derived carbon, disturbances in plant cover are thought to be detrimental for the persistence of soil biota. In this work, we studied the disturbance effects of plant removal and soil mixing and the mitigation effects of replanting on soil biota in a low-arctic meadow ecosystem. We set up altogether six replicate blocks, each including three randomized treatment plots, at two distinct fells at Kilpisjärvi, northern Finland. Vegetation was removed in two thirds of the plots: one third was then kept barren (the plant-removal treatment), while the other third was replanted with a local herb Solidago virgaurea. The remaining plots of intact vegetation were used as treatment comparisons. The responses of soil microbes and fauna were examined six years later in the early and late growing season. The biomass of bacteria, non-mycorrhizal fungi and mycorrhizal fungi (estimated using PLFA markers) were on average 74%, 89% and 84% lower in the plant-removal and 64%, 74% and 71% lower in the Solidago replant plots than in the intact meadow. The positive effect of replanting was statistically significant for fungi, but not for bacteria. The PCA of relative PLFA concentrations further showed that the structure of the microbial community differed significantly among all three treatments. The abundance of nematodes and collembolans was on average 82 and 95% lower, but the total number of nematode genera and collembolan taxa only 27 and 7% lower in the plant-removal plots than in the intact meadow soil. Few disturbance effects on soil fauna were significantly mitigated by the Solidago replant (the plant parasitic nematodes being a notable exception) and in the case of the collembolans, the Solidago replant plots had even fewer animals than the plant-removal plots. The response of soil biota also varied with locality: the effects on fungivorous nematodes were found at one site only and the replant effects on the number and diversity of collembolan taxa varied with site. Our results suggest that despite drastic reductions in the abundance of soil biota, the majority of animal taxa can persist for years in disturbed arctic soils in the absence of vegetation. In contrast, the alleviating replant effects on the abundance of soil biota appear weak and may only partially reverse the negative effects of vegetation removal and soil disturbance.     

Influence of erosion on soil microbial biomass,abundance and community diversity

This study aimed to determine microbial biomass carbon and microbial abundance immediately after, and two years after, forest soil erosion, so as to estimate the degree of damage, including the rate of recovery of microorganisms, in each area. It also aimed to determine the community diversity, and to establish relationships between microbial biomass, microbial abundance and the physico‐chemical properties of the soil. Three different study areas in Hiroshima Prefecture, Japan, were used. One undisturbed area and two eroded areas (one immediately after and one two years after erosion). The analysis of variance showed a highly significant difference in microbial biomass carbon and abundance between the study areas. The undisturbed area showed the highest value, followed by the area eroded two years ago, then lastly the area studied immediately after the erosion. The biomass carbon was highly correlated with gram positive bacteria with r² = 0·983, p < 0·01. The biomass carbon and microbial population were shown to be significantly correlated to the soil's physico‐chemical properties, such as pH, moisture content, water‐holding capacity and CN ratio. However, CN ratio proved to be closely correlated to biomass carbon with r² = −0·978, p < 0·01, to Gram‐positive bacteria with r² = −0·977, p < 0·01, to Gram‐negative bacteria with r² = −0·989, p < 0·01 and to fungi with r² = −0·977, p < 0·01. The undisturbed area showed a highly diverse community in both of the restriction enzymes used, followed by the area affected by erosion two years ago, then the area immediately after erosion. Copyright © 2004 John Wiley & Sons, Ltd.     

Effects of compaction on soil surface water repellency

Water repellency can reduce the infiltration capacity of soils over timescales similar to those of precipitation events. Compaction can also reduce infiltration capacity by decreasing soil hydraulic conductivity, but the effect of compaction on soil water repellency is unknown. This study explores the effect of compaction on the wettability of water repellent soil. Three air‐dry (water content ～4 g 100 g^?1) silt loam samples of contrasting wettability (non‐repellent, strongly and severely water repellent) were homogenized and subjected to various pressures in the range 0–1570 kPa in an odeometer for 24 h. Following removal, sample surface water repellency was reassessed using the water drop penetration time method and surface roughness using white light interferometry. An increase in compaction pressure caused a significant reduction in soil surface water repellency, which in turn increases the soil's initial infiltration capacity. The difference in surface roughness of soils compacted at the lowest and highest pressures was significant (at P > 0.2) suggesting an increase in the contact area between sessile water drops and soil surfaces was providing increased opportunities for surface wetting mechanisms to proceed. This suggests that compaction of a water repellent soil may lead to an increased rate of surface wetting, which is a precursor to successful infiltration of water into bulk soil. Although there may be a reduction in soil conductivity upon compaction, the more rapid initiation of infiltration may, in some circumstances, lead to an overall increase in the proportion of rain or irrigation water infiltrating water repellent soil, rather than contributing to surface run‐off or evaporation.     

Effects of pre-sowing compaction on soil physical properties, soil atmosphere and growth of oats on a clay soil

The effects on a number of soil physical and aeration parameters of compaction during spring pre-sowing operations were measured on a clay soil (49% clay). A soil-tyre contact stress of 200 kPa was applied by tractor tyres.
Yield of an oat crop was reduced by 30% as a result of compaction. Total porosity of the soil was reduced by 6% v/v owing to loss of pores > 60 μm, and water retention was increased. The resultant decrease in air-filled porosity greatly reduced gas diffusion and air permeability coefficients of the soil, and, for a time, O₂ content of the soil atmosphere was significantly lowered in the compacted treatment. Penetrometer resistance after sowing was 3.5 MPa in the control and 4.5 MPa in the compacted treatment; in the latter, root growth was inhibited until the soil dried and cracked. By the end of June, canopy temperature measurements indicated water stress in the oat crop on compacted soil but not in that on the control.
The results obtained indicated that air permeability, measured in the field, of 1 mm s⁻¹ provides a satisfactory single value below which crop growth is likely to be reduced.     

Effects of soil compaction on the relationships between nematodes,grass production and soil physical properties

As farm machinery has become heavier, concern has grown about its direct effects on soil physical conditions and its indirect effects on crop yields and soil biota. To study the relationships between these parameters, non-grazed temporary grassland plots on a loamy sand soil were subjected to full-width load traffic with widely different loads (0, 4.5, 8.5 and 14.5 t) one to four times per year for a period of 5 years. Soil bulk density was monitored as an indicator of soil compaction. Grass yield was measured throughout the experimental period. Root distribution over the soil profile and nematodes populations were assessed during the final year of the experiment. Results indicate that a moderate degree of compaction (∼4.5 t load) gave the highest crop yield and that at higher degrees of compaction roots failed to penetrate into the deeper soil layers (>20 cm depth). Total numbers of nematodes were not affected by compaction, but their distribution over the various feeding types shifted towards a population with increased numbers of herbivores and decreased numbers of bacterivores and omnivores/predators. This change in the structure of the nematode assemblage is associated with poorer conditions for crop growth.     

土壤压实对土壤物理性质及小麦氮磷钾吸收的影响

为了研究土壤压实对土壤物理性质以及小麦养分吸收情况的影响,在2006和2007年进行了两轮田间试验.试验中,先用旋耕机对田块进行旋耕,耕深10cm,然后使用手扶式、轮式、履带式拖拉机在旋耕后的田块中通过1次(T1)、2次(T2)、4次(T3)以对土壤进行压实处理,对照组(T4)不作任何压实处理.压实处理后再次对土壤表层进行浅旋耕,耕深5 cm,耕后用播种机进行小麦播种,小麦品种为南京-601.试验结果发现,次表层土壤的压实处理显著影响次表层土壤的容重,孔隙度,小麦蛋白质含量以及植物中N、P、K的含量.除次表层的土壤容重在T3组中最大,T4组中最小外,其他参数值在T4组中最大,T3组中最小.并且,随着次表层土壤压实程度的增加,几乎所有的参数(土壤容重除外)都有所减少.不过,与第一年相比,参数值在第二年略有增加.总之,土壤压实严重破坏土壤结构,不利于小麦对养分的吸收.     

Effects of organic matter removal, soil compaction, and vegetation control on Collembolan populations

Collembola can be among the most numerous meso-invertebrates in the forest floor and, through their interaction with primary decomposers in the decomposition food web, may affect litter decomposition and consequently site productivity. This study was conducted to determine whether Collembolan abundance could be impacted by organic matter removal, compaction, and vegetation control on a loblolly pine (Pinus taeda L.) plantation. Monthly soil and litter samples were taken over 2 years and the fauna extracted from them using modified Tulgren funnels. Organic matter removal and vegetation control generally caused a significant decrease in Collembolan populations, while compaction did not significantly affect Collembolan populations. These results indicate that habitat was the primary influence on population abundance in this experiment, possibly via its influence on desiccation. Sensitivity of collembolan populations to habitat changes caused by organic matter removal indicates a potential effect on long-term site productivity.     

Drilling vegetables into autumn-cultivated soil with a low ground pressure vehicle: effects on timeliness and soil compaction

Secondary cultivations for the spring-sowing of vegetables are often delayed because they cannot be carried out until the soil to the depth of cultivation has dried to the plastic limit (i.e. it crumbles rather than smears). It is suggested that doing secondary cultivations in the autumn, and then drilling in the spring without further cultivation, would be less liable to delays because only the soil down to the depth of drilling need dry below the plastic limit. Simple models are derived to calculate soil water contents from weather data. They show that between 1960 and 1981 the 0–120 mm layer (representing a typical cultivation depth) dried beyond the plastic limit for an average of 36% of the days between 14 February and 9 May, compared with 68% for the 0–30 mm layer (proposed system). Wheel sinkage, bulk density and cone resistance measurements showed that a tractor would cause excessive compaction when only the surface 30 mm was drier than the plastic limit, but this could be minimized by using a low ground pressure vehicle.     

大中小型拖拉机压实对土壤坚实度和大豆产量的影响

探讨农业机械压实对土壤坚实度和产量的影响规律,对改善作物生产环境、促进农业机械化向质量型转变具有重要意义。以东北典型黑土区耕地土壤为研究对象,依照随机区组试验原理,选择大、中、小3种型号拖拉机进行6种压实处理,同型拖拉机相同压实次数试验重复3次,采用PV6.08型贯穿阻力仪测量压实轮辙截面土壤坚实度。试验结果表明：土壤坚实度随压实次数增加而逐渐递增,3种拖拉机压实测试截面浅层均出现明显压实核,且压实核内土壤坚实度随压实次数增加而逐渐增大,CASE-210型拖拉机压实对表层土壤坚实度影响程度和范围最大,压实12次时压实核处土壤坚实度达4.0 MPa,JD-280型拖拉机对深层土壤压实影响程度和范围最大,在65~80 cm的土壤深层坚实度的峰值达3.2 MPa;拖拉机压实均导致大豆产量降低,CASE-210、JD-904和JD-280拖拉机压实12次时大豆产量分别降低了21.24%、18.15%和12.38%。     

土壤紧实胁迫对黄瓜碳水化合物代谢的影响

用容重分别为1.25 g/cm3(疏松土壤,即对照)和1.55 g/cm3(紧实土壤)的土壤进行盆栽试验,研究了土壤紧实胁迫对“津春4号”黄瓜(Cucumis sativusL.)不同生育期叶片和根系碳水化合物代谢的影响,以探讨土壤紧实胁迫对黄瓜生长产生影响的机理.结果表明,在土壤紧实胁迫条件下,黄瓜不同生育期叶片的净光合速率(Pn)、气孔导度(Gs)和蒸腾速率(Tr)均显著下降,胞间CO2浓度(Ci)显著升高,光合作用受到抑制;叶片中蔗糖磷酸合成酶(SPS)活性显著降低,蔗糖合成酶(SS)、酸性转化酶(AI)和中性转化酶(NI)活性显著增强,蔗糖、葡萄糖、果糖和淀粉含量显著增加,蔗糖的合成与输出受到抑制;不同生育期根系SPS、AI和NI活性显著下降,而SS活性显著增强,蔗糖、葡萄糖和果糖含量显著增加,淀粉含量基本不变.这表明,土壤紧实胁迫抑制了黄瓜叶片中同化物的合成和输出,降低了碳水化合物向根系中的输入,阻碍了根系对碳水化合物的利用,使植株矮小,产量下降.     

土壤紧实胁迫对黄瓜生长、产量及养分吸收的影响

用容重分别为1.2、1.4和1.6.g/cm3的土壤进行盆栽试验,研究了土壤紧实度对黄瓜生长、产量及养分吸收的影响。结果表明,当土壤紧实度增大时,黄瓜秧苗的株高在定植后的15.d后受到显著抑制;第4叶的叶宽和叶长在定植后9～17.d内增加;茎粗则是在稍紧的土壤中(R.1.4)最大,过紧的土壤中(R.1.6)最小;根系伸长生长受阻,干物质质量及活力显著下降,根冠比降低;生物学产量、经济产量、经济系数的变化情况及植株对氮、磷、钾吸收量的变化与茎粗的变化趋势相同。在本试验条件下,容重为1.2.g/cm3的土壤利于株高及根系的生长,容重1.4g/cm3的土壤则利于茎粗、根系养分的吸收及产量的增加。     

Influence of disturbance and nitrogen addition on plant and soil animal diversity in grassland

Two key determinants of biological diversity that have been examined in aboveground and aquatic systems are productivity, or resource supply, and physical disturbance. In this study, we examined how these factors interact under field conditions to determine belowground diversity using microarthropods (mites and Collembola) as our test community. To do this, we established a field manipulation experiment consisting of crossed, continuous gradients of nitrogenous (N) fertilizer addition (up to 240 kg N ha^?1) and disturbance (imitated trampling by cattle) to produce a gradient of soil nutrient availability and disturbance. Due to the relatively short-term nature of our study (i.e. 2 years), we only detected minimal changes in plant diversity due to the experimental manipulations; in the longer term we would expect to detect changes in plant diversity that could potentially impact on soil fauna. However, disturbance reduced, and additions of N increased, aboveground biomass, reflecting the potential effects of these manipulations on resource availability for soil fauna. We found that disturbance strongly reduced the abundance, diversity, and species richness of oribatid mites and Collembola, but had little effect on predatory mites (Mesostigmata). In contrast, N addition, and therefore resource availability, had little effect on microarthropod community structure, but did increase mesostigmatan mite richness and collembolan abundance at high levels of disturbance. Oribatid community structure was mostly influenced by disturbance, whereas collembolan and mesostigmatan diversity were responsive to N addition, suggesting bottom-up control. That maximal species richness of microarthropod groups overall occurred in undisturbed plots, suggests that the microarthropod community was negatively affected by disturbance. We found no change in microarthropod species richness with high N additions, where plant productivity was greatest, indicating that soil biotic communities are unlikely to be strongly regulated by competition. We conclude that the diversity of soil animals is best explained as a combination of their many varied life history tactics, phenology and the heterogeneity of soils that enable so many species to co-exist.     

Effects of aggregate size,soil compaction,and bovine urine on N2O emissions from a pasture soil

The dominant N₂O emission source in New Zealand, calculated using the Intergovernmental Panel on Climate Change methodology, is agricultural soils. The largest source of N₂O emissions in New Zealand occurs as a result of excreta deposition onto pasture during grazing. There is a dearth of studies examining the effect of soil compaction and soil aggregate size on N₂O emissions from urine patches in grazed pastures. In this study, we repacked soil cores with four different soil aggregate sizes (<1.0–5.6 mm diameter), applied bovine urine, and then subjected the soil cores to four levels of soil compaction. Fluxes of N₂O were monitored for 37 days after which soil cores were allowed to dry out prior to a rewetting event. There was an interaction between aggregate size and soil compaction with the cumulative loss of N₂O over the first 37 days ranging from 0.3% to 9.6% of the urine-N applied. The highest N₂O emissions occurred from the smallest and most compacted aggregates. Even under the highest levels of compaction the N₂O loss from the large aggregates (4.0–5.6 mm diameter) was <1% of the urine-N applied. Reasons for the observed differences in the N₂O flux from the different-sized aggregates included varying gas diffusivities and higher rates of denitrification in the smallest aggregates, as evidenced by the disappearance of nitrate.     

SSA土壤固化剂对黄土击实、抗剪及渗透特性的影响

为了探讨土壤固化剂对黄土力学特性的影响,通过黄土样中加入SSA土壤固化剂前后的击实试验、直剪试验和渗透试验的对比分析,研究了固化剂掺量、养护龄期与固化土击实、抗剪强度及渗透特性的关系。结果表明,随SSA土壤固化剂掺量增加,固化土的最优含水率有所降低,最大干密度有所增大;固化土的抗剪强度指标黏聚力和内摩擦角随固化剂掺量的增加和养护龄期的延长而增大,渗透系数随固化剂掺量的增加和养护龄期的延长而减小。在实际应用中,建议SSA固化剂最佳掺量为1%,养护龄期至少7 d以上。     

Effects of foliar and soil insecticide applications on the collembolan community of an early set-aside arable field

Effects of foliar and soil insecticide applications on collembolan density and community structure were investigated in an early set-aside arable field. Insecticides were applied separately and in combination to the soil surface (chlorpyrifos) and vegetation (dimethoate). The treatments were established to investigate effects of above- and below-ground insects on plant succession. Starting in 1997, the insecticides were applied from April to November at 2-week (dimethoate) or monthly intervals (chlorpyrifos). Samples were taken in 2000 prior to and after insecticide application in March and June, respectively. Both insecticides are lethal to Collembola and insecticide applications resulted in a strong decline in the density of total Collembola. Application of chlorpyrifos reduced collembolan density to a greater extent than dimethoate; the effect of the combined application on total collembolan numbers was similar to that of chlorpyrifos only. Collembolan numbers recovered after the insecticide applications in 1999, but in the treated plots populations were still reduced in March 2000 before the re-application of insecticide treatments in that year. The insecticide applications changed the dominance structure of the collembolan community, but had no effects on species composition. The results may be of relevance for the interpretation of studies on plant–insect herbivore interactions using insecticides.     

Effects of biochar,earthworms, and litter addition on soil microbial activity and abundance in a temperate agricultural soil

Biochar application to arable soils could be effective for soil C sequestration and mitigation of greenhouse gas (GHG) emissions. Soil microorganisms and fauna are the major contributors to GHG emissions from soil, but their interactions with biochar are poorly understood. We investigated the effects of biochar and its interaction with earthworms on soil microbial activity, abundance, and community composition in an incubation experiment with an arable soil with and without N-rich litter addition. After 37 days of incubation, biochar significantly reduced CO₂ (up to 43 %) and N₂O (up to 42 %), as well as NH₄ ⁺-N and NO₃ ^?-N concentrations, compared to the control soils. Concurrently, in the treatments with litter, biochar increased microbial biomass and the soil microbial community composition shifted to higher fungal-to-bacterial ratios. Without litter, all microbial groups were positively affected by biochar × earthworm interactions suggesting better living conditions for soil microorganisms in biochar-containing cast aggregates after the earthworm gut passage. However, assimilation of biochar-C by earthworms was negligible, indicating no direct benefit for the earthworms from biochar uptake. Biochar strongly reduced the metabolic quotient qCO₂ and suppressed the degradation of native SOC, resulting in large negative priming effects (up to 68 %). We conclude that the biochar amendment altered microbial activity, abundance, and community composition, inducing a more efficient microbial community with reduced emissions of CO₂ and N₂O. Earthworms affected soil microorganisms only in the presence of biochar, highlighting the need for further research on the interactions of biochar with soil fauna.     

The effect of soil compaction, profile disturbance and fertilizer application on the growth of eucalypt seedlings in two glasshouse studies

Soil damage, compaction and displacement, during logging or clearing and cultivation affects both soil physical and chemical properties and reduces growth of regenerated or planted tree seedlings. Understanding the factors involved will aid management and set limits for indicators of sustainable management in eucalypt forests. In the first of two glasshouse studies, three Eucalyptus species were grown for 110 days in soils from six forest sites in Tasmania, Australia. Sites sampled ranged from low rainfall dry forest to very high rainfall wet forest. Soil was collected from three soil depths, in 10 cm increments to 30 cm, each packed in pots to four different bulk densities, ranging from that present in undisturbed field sites to that plus 0.17 g cm⁻³. In the second study Eucalyptus globulus Labill. seedlings were grown in soil collected from disturbed and undisturbed sites, packed to two bulk densities, and fertilized with combinations of N and P. Increasing soil compaction, in Study 1, caused a proportional decrease in final mass of seedlings of up to 25%. Growth on soil from lower horizons (10–30 cm) averaged only 41% of that on topsoil, a significantly greater restriction of growth than that achieved through compaction. It was concluded that topsoil displacement and profile disturbance was a more significant form of soil damage than compaction. Above-ground dry weight of seedlings was most strongly correlated with soil total N but poorly correlated with other macronutrients. Growth of E. globulus seedlings grown on disturbed soils, in Study 2, averaged 30% of that on undisturbed sites. With added P and N on undisturbed sites growth averaged seven times that of the unfertilized seedlings indicating a general deficit of available P and N on the three soils tested. On soils from disturbed areas, there was also a response to fertilizing with N and P together but the response varied on the three soils. The effects of profile disturbance were ameliorated with fertilizer applications on only one of the soils. The results highlighted the importance of retaining topsoil in situ during forest operations.     

Effects of compaction on soil hydraulic properties,penetration resistance and water flow patterns at the soil profile scale

The aim of this study was to quantify the effects of compaction on water flow patterns at the soil profile scale. Control and trafficked plots were established in field trials at two sites. The trafficked treatment was created by four passes track‐by‐track with a three‐axle dumper with a maximum wheel load of 5.8 Mg. One year later, dye‐tracing experiments were performed and several soil mechanical, physical and hydraulic properties were measured to help explain the dye patterns. Penetration resistance was measured to 50 cm depth, with saturated hydraulic conductivity (K_s), bulk density, and macroporosity and mesoporosity being measured on undisturbed soil cores sampled from three depths (10, 30 and 50 cm). Significant effects of the traffic treatment on the structural pore space were found at 30 cm depth for large mesopores (0.3–0.06 mm diameter), but not small mesopores (0.06–0.03 mm) or macroporosity (pores > 0.3 mm). At one of the sites, ponding was observed during the dye‐tracing experiments, especially in the trafficked plots, because of the presence of a compacted layer at plough depth characterized by a larger bulk density and smaller structural porosity and K_s values. Ponding did not induce any preferential transport of the dye solution into the subsoil at this site. In contrast, despite the presence of a compacted layer at 25–30 cm depth, a better developed structural porosity in the subsoil was noted at the other site which allowed preferential flow to reach to at least 1 m depth in both treatments.     

Measurement, interpretation and modelling of soil compaction

Abstract. The analysis of some experimental field results is used to illustrate the problem of measuring and evaluating compaction treatment effects below wheel ruts of different depths. A solution to the problem is described which traces vertical soil movement to allow comparisons between treatments to be made using soil elements which derive from the same depth in the undisturbed profile, irrespective of their depths in the compacted profile.
A soil compaction model, which predicts the changes in dry bulk density resulting from the passage of wheels, is briefly described. Examples are given of its use in comparing the compaction caused by various types and arrangements of wheels and in assessing the contribution made by a particular input variable.