Abundance of biogenic structures of earthworms and termites in a mango orchard

A comparative study of the spatial distributions and the quantity of biogenic structures produced by earthworms and termites (Odontotermes nilensis and Ancistrotermes guineensis) has been conducted in a mango orchard at Thiès (Senegal).This study showed that surface biogenic structures may represent a large amount of modified soil (up to 536.5 g m⁻²) which vary depending upon the seasons and the species. Whilst the quantity of casts was independent on the season (178.6 g m⁻²), O. nilensis sheetings fluctuated with the seasons. In addition, we show that the spatial organisation of surface biogenic structures fluctuates with seasons. It displays patches ranging from 5 to 15 m. There is a link between the distribution of earthworm casts and the vegetation. In addition, spatial distribution was also linked to the biology of constructing species. We observed that the A. guineensis’ filling structures were mainly located under the mango trees during the dry season where the stems and the brushwoods were abundant. It appears that the spatio-temporal distribution of the biogenic structures under study depended upon two main factors: season and vegetation. However, depending upon the biology of the engineer, these two factors influenced the spatial distribution of structures in different ways.     

Accumulation of heavy metals by enchytraeids and earthworms in a floodplain

The river floodplain ‘Afferdense and Deestsche Waarden’ (ADW) in The Netherlands is diffusely contaminated with several heavy metals. It is, however, unclear whether this mixed contamination exerts any adverse ecotoxicological effects. In November 2000 and May 2001 a field survey was conducted in two areas in the ADW to collect a wide range of data concerning contamination levels, bioavailability, enchytraeids and earthworms and abiotic factors such as lutum and organic matter content, cation exchange capacity (CEC) and soil nutrient concentrations. Earthworms and enchytraeids were also analyzed for heavy metal content. At both sites arsenic and zinc were present in soil at relatively high concentrations (above the Dutch intervention value). In the two areas, both enchytraeids and earthworms accumulated metals. Fridericia ulrikae accumulated more cadmium than Enchytraeus buchholzi and Henlea perpusilla. The earthworm Lumbricus rubellus accumulated larger concentrations of Cr, Cu and Pb than Aporrectodea caliginosa and Allolobophora chlorotica. Dietary, physiological and behavioral characteristics may have contributed to these differences.     

The influence of livestock dung on the abundance of exotic and native earthworms in a grassland in south-eastern Australia

It is well known that organic matter in the form of dung is utilised as a food source by some earthworms, but little has been reported on the preferences of earthworms for different types of dung in agricultural settings. An experiment was carried out in spring in south-eastern Australia to evaluate the effect that dung from different livestock has on the abundance of earthworms in a grassland environment. We were particularly interested to compare the responses of native Australian earthworms (Megascolecidae) with those of exotic earthworms (Lumbricidae and Acanthodrilidae). The attractiveness of dung from sheep, cattle and horses was measured by determining the abundance and biomass of the resident earthworm species under each dung type at varying times after adding the dung to the soil surface (0, 10, 20 and 30 days). The earthworm population consisted of three exotic species, Apporrectodea trapezoides, Microscolex dubius and M. phosphoreus, and two native species, Spenceriella macleayi and S. bywongensis. Both the number of days that the dung was available to earthworms and the type of dung influenced the numbers and biomass of the earthworms found beneath the dung pats. Significant interactions existed between time and dung type when all adult earthworms were considered as one group and also when adults were split into individual species. The various species responded differently to the dung, but horse dung was generally the more preferred dung type. The significance of these results is discussed in terms of the management of dung in an Australian pastoral context.     

Habitat use and activity patterns of larval and adult Cantharis beetles in arable land

Generalist predators play a key role in agriculturally and environmentally sustainable systems of pest control. A detailed knowledge on their ecology, however, is needed to improve management practices to maximize their service of pest control. The present study examines the habitat use and activity patterns of larval and adult Cantharis beetles that are abundant predators in arable land. Laboratory experiments revealed that sixth instar larvae of Cantharis fusca and Cantharis livida significantly preferred high relative humidity levels of 85–90% to lower ones. This can explain their preference for meadows over fields due to the more favorable microclimatic conditions in the former habitats. Surface activity of sixth instar Cantharis larvae during autumn, winter and early spring occurred at soil temperatures above 0 °C. However, no correlation between surface activity and soil temperature, air temperature or relative humidity was found above 0 °C. Catches of sixth instar Cantharis larvae within fenced pitfall traps were higher in a meadow (Mean ± S.D.; 13.8 ± 7.63 individuals m⁻²) than in a field (4.60 ± 2.89 individuals m⁻²). Mark-recapture density estimations for sixth instar larvae indicated mean densities of 25.9 ± 5.63 (field) and 42.8 ± 16.0 individuals m⁻² (meadow). The same pattern was found for adult emergence rates in the field (0.17 ± 0.39 adults m⁻²) and meadow (1.83 ± 1.17 adults m⁻²) as well as for adult densities in the vegetation (field 4.89 ± 3.62 adults 60 m⁻²; meadow 12.5 ± 11.2 adults 60 m⁻²). It is concluded that especially in winter elements that provide plant cover should be incorporated in arable fields to enhance larval cantharid population densities and to attract them from their prime grassland habitats into arable sites.     

The efficiency of soil hand-sorting in assessing the abundance and biomass of earthworm communities. Its usefulness in population dynamics and cohort analysis studies

Pit digging and manually revising soil blocks is a frequently used method used for field studies of earthworm communities. The aim of this study was to compare the efficiency of hand-sorting (HS) to extract small earthworms, ca. 0.2 g, and the usefulness in studies of population dynamics and cohort analysis. Many earthworms are not recovered when revising manually the soil. Factors include soil characteristics, i.e. moisture, texture, etc. and also a human factor, which is more relevant if the study is conducted in the long-term. We used data collected in a field study of earthworm communities during 2 years in the savannas of Colombia. Small soil blocks (20 × 20 × 20 cm) were dug out in order to collect the smallest earthworms by washing-sieving (WS) and compare the results with the standard HS of large monoliths (100 × 100 × 50 cm). In fact, this methodology has rarely been addressed in earthworm population field studies. Our results showed that HS efficiency varied owing to the species and ranged from 31.4% up to 100% in the savanna and from 44% to 80% in the pasture, for two small species, i.e Aymara n. sp. (epigeic) and Ocnerodrilidae sp. (endogeic). In the case of the Glossodrilus n. sp. (endogeic) these values were similar, i.e. 51.7% and 58.1%, in the savanna and pasture, respectively. We also used frequency tables to calculate the average efficiency of HS 1 m² soil cores for each weight class in each species in order to obtain a population density correction factor. This allowed us to make corrections in earthworm density in the histograms for population dynamics analysis. We conclude that this method should be the modus operandi in long-term earthworm demography studies.     

Effects of disturbance on the biodiversity and abundance of isopods in temperate grasslands

Connell's ‘intermediate disturbance hypothesis’ was tested using a relatively simple, species poor isopod community in grass heaths and in a coastal sand dune grassland. Variation in diversity resulted from disturbance both to the soil profile and sward structure due to grazing. Predictions from a spatial survey were tested using a 20 years time series. These changes were related to changes in rabbit activity, anthropogenic soil disturbance and on one site, deposition of blown sand. Changes in the soil profile resulted in diversity initially increasing due to decreases in the density of the commonest species, leading to an increase in equitability. Prolonged disturbance to the soil profile both due to sand blowing onto one site and rotavating on another, lead to a reduction in diversity due to one of the species being eliminated. The isopod community was most diverse when least disturbed by grazing, of intermediate diversity under intermediate grazing intensity and least diverse when most heavily grazed. In these simple communities, differences in diversity were mostly determined by differences in equitability rather than in species richness. We conclude that maximising heterogeneity of sward structure could have advantages for the conservation of both invertebrate biodiversity and vertebrates that feed on them.     

Long-term effects of leguminous cover crops on biochemical and biological properties in the organic and mineral layers of soils of a coconut plantation

The primary aim of the study was to determine the long-term (12 years) effects of leguminous cover crops like Atylosia scarabaeoides, Centrosema pubescens, Calopogonium mucunoides and Pueraria phaseoloides on important soil biochemical and biological properties and their interrelationships in the organic (fresh litter layer, F and fermented + humus layer, F + H) and mineral (0–10 and 10–20 cm) layers of soils of a 19-year-old coconut plantation.The total biomass production (above-ground) for the 12-year period varied significantly between the cover crops and ranged from 34.86 (calopo) to 90.43 (pueraria) Mg ha^–1. Total N and C additions at the cover cropped (CC) site for the 12-year period were 0.97–3.07 Mg ha^–1 and 16.90–43.34 Mg ha^–1, respectively. Irrespective of layers, the levels of organic C, total N, organic substrates viz., dissolved organic C and N, labile organic N, water soluble carbohydrates, and light fraction organic matter-C and were markedly higher in the CC site compared to the control. Consequently, the levels of microbial biomass-C (C_MIC), -N (N_MIC) and -P (P_MIC), net N mineralization rates, CO₂ evolution, metabolic quotient (qCO₂) and the activities of l-asparaginase, l-glutaminase and β-glucosaminidase were significantly higher in the CC site compared to the corresponding levels in the control site. Between layers, the levels of various chemical, biochemical and microbial parameters were consistently higher in the organic layers compared to the mineral layers at all the sites including control. Among the ratios of various microbial indices, the ratios of C_MIC: organic C and C_MIC: P_MIC did not differ significantly between the layers and sites. However, the ratio of C_MIC: N_MIC was relatively higher in the mineral layers and control site. The variation in individual soil properties between layers and sites reflected the concomitant changes occurring in soil organic matter content. Apparently, microbial activity was limited by the supply of biologically available substrates in the mineral layers and the control site. Contrarily, the more direct supply of nutrients from decomposing plant litter and the indirect supply of nutrients from the mineralization of organic matter led to significantly higher levels of microbial biomass in the organic layers.     

Biodiversity and abundance of terrestrial isopods along a gradient of disturbance in Sabah,East Malaysia

Connell's intermediate disturbance hypothesis predicts that the highest diversity is maintained at intermediate levels of disturbance. We have examined this hypothesis by observing differences in biodiversity of terrestrial isopods along a gradient of disturbance from two undisturbed primary tropical rainforest sites, to a logged site, a mixed native fruit orchard and a commercial oil palm plantation, in Sabah, East Malaysia. We describe a standardised protocol for the rapid assessment of isopod biodiversity on tropical forest floor sites and for measuring environmental variables to which we have related differences in species richness and relative abundance of the isopods. The results do not support Connell's hypothesis because there were no significant differences in diversity, species richness or equitability between disturbed sites and the nearest primary forests. The relative abundance of individual species was highest in the most disturbed environment. We suggest that this may be because particular species are well adapted to exploiting resources under the more ‘r’ selection conditions created by disturbance. Possible reasons for why the observations do not conform with predictions from the intermediate disturbance hypothesis are discussed. We conclude that Huston's dynamic equilibrium model is more appropriate than the intermediate disturbance hypothesis in predicting the effects of disturbance of tropical rainforests on these arthropod macro-decomposers.     

Indirect gradient analysis at different spatial scales of prorated and non-prorated earthworm abundance and biomass data in temperate agro-ecosystems

We investigated how earthworm communities of agricultural systems vary in abundance, biomass and species composition at different spatial scales. In four farms representative of the main agro-ecological regions of Belgium 14 parcels were sampled using a combined method (hand sorting after formalin extraction). Parcels in both grassland and arable land were studied. In each parcel two or three sample plots (1 m²) were randomly selected; these were further split up in two, resulting in four or six 0.5 m² subplots per parcel. Principal component analysis was applied to earthworm numbers and biomass. Innovative was that we compared the results from data matrices with raw data with prorated matrices, where unidentified juveniles were added to species following certain rules. To test the degree of variability between the different spatial levels, average distances between subplots, plots and parcels based on ordination scores were compared. A general linear model with a nested structure for subplots, plots, and parcels was built to find absolute differences between farms. Earthworm numbers ranged from 10 to 463 individuals m^–2, biomass varied between 3 and 186 g m^–2 and species numbers ranged from one to seven per parcel. Ordinations separated earthworm communities along a geographical—mainly soil—gradient on the first axis and distinguished between land uses along the second axis. Both in numbers of individuals and in biomass differences in earthworm populations increased from subplot to parcel level. As expected, differences within a pair of subplots were small. Plots within the same parcel tended to be highly similar, but in some cases large dissimilarities were encountered, reflecting a clustered population structure of earthworm communities at the field scale, and, possibly, abiotic gradients. The largest differences were observed at the parcel level. The latter may be attributed to differences in land use. Prorating had a narrowing effect on the data through which differences became less pronounced.     

Response of soil micro- and mesofauna to diversity and quality of plant litter

The abundance and functional structure of soil micro- (nematodes) and mesofauna (collembolans and mites) in relation to species diversity and initial C:N ratio of plant litter were studied in a field mesocosm experiment. A total of five litter treatments were applied to generate an increasing diversity of plant species (one, three and 12 species) and/or differences in initial C:N ratio of the litter (low, intermediate and high ratio). Samples were taken 3, 6 and 24 months after the litter exposure. On each sampling date litter and underlying sand samples were taken. Our results showed that litter quality, but not litter diversity was the factor which affected the three animal groups under study. The effect of litter was dependent on the time of litter exposure. Nematode fauna colonized litter earlier than the two mesofaunal groups. Nematodes responded apparently to litter quality gradient at early stages of litter decomposition. Three months after the start of the experiment the highest density of nematodes was noticed in single species litter of Trifolium pratense. Bacterial-feeding nematodes dominated in all litter treatments; on the first sampling date their percent share in Trifolium litter reached even 99.9%. Opposite to nematodes at late stages of litter decomposition the two mesofaunal groups seemed to show some preferences for low quality litter of Festuca rubra. Collembolan and nematode diversity was affected in similar way by the litter quality; the lowest diversity of the animal communities was found in the litter of the lowest initial C:N ratio. Maturity index of nematode communities was found to be a good index to differentiate between litters of different quality. The abundance and community structure of the three animal groups in underlying sandy soil was not significantly influenced by experimental conditions.     

Effects of Cd and Pb on the survival and juvenile production of Sinella coeca and Folsomia candida

Degradation factors can cause both quantitative and qualitative changes in fauna and thus affect soil functioning. The use of bio-indicators may help to detect environmental changes. The aim of this study was to evaluate the effects of lead and cadmium on the biology (mortality, reproduction, egg hatching) of two euedaphic collembolans, Sinella coeca and Folsomia candida and to evaluate sensitivity differences between the two species. The results showed that low concentrations of lead did not cause dramatic effects on the survival and reproduction of both species. Reproduction of both species was reduced when the soil contained a nominal concentration of Pb at 1000 μg g^–1. S. coeca and F. candida showed a high sensitivity to cadmium and both their reproduction and survival were significantly reduced on exposure to 50 μg g^–1. Moreover, egg hatching of F. candida was affected by the cadmium concentrations in the soil, and by the amount of time that the eggs stayed in contact with cadmium. This study showed that the inhibition determined by Cd on F. candida juvenile production was caused in part by inhibition of deposition and, in part, by reduction of egg viability.     

Long-term consequences of biochemical and biogeochemical changes in the Horseshoe Bend agroecosystem,Athens, GA

Understanding many soil processes, including the accumulation of organic matter and the formation and loss of soil aggregates, requires research that is conducted over decadal time periods. The dynamics of soil organic matter and soil fauna at the Horseshoe Bend (HSB) agroecosystem site in Georgia have been studied in replicated experimental plots since 1978. The experimental treatments (no-tillage (NT) and conventional-tillage (CT) regimes) are continuing to diverge in amounts and distribution of SOM in the soil profiles of HSB. Our current research focuses on two major areas: 1) long-term measurements of the gradually-increasing base of soil organic matter from C3-pathway plants, in crop rotations that have been in effect since 1997; 2) following the production, accumulation, fate and ecological effects of the Bt (Bacillus thuringiensis) proteins from the summer planting of Bt (and non-Bt) cotton in subplots within our main plots. The variation in the size of soil aggregates may influence the sequestration of Bt toxins, and their breakdown products, within soils. NT management systems at HSB generate an increasing proportion of soil macroaggregates in comparison to the markedly reduced macroaggregates in our CT plots. We suggest that NT systems may sequester more Bt-related products than will CT plots.     

Effects of carbendazim and lambda-cyhalothrin on soil invertebrates and leaf litter decomposition in semi-field and field tests under tropical conditions (Amazônia,Brazil)

Effects of the fungicide carbendazim and of the insecticide lambda-cyhalothrin on soil invertebrates and litter decomposition under tropical conditions were assessed in ecotoxicological semi-field studies using intact soil-core terrestrial model ecosystems (TMEs) and in a field test applying the litter-bag method. In the TME study, performed indoors under controlled conditions, earthworms, isopods and diplopods were added to intact soil cores and mortality of soil invertebrates and mass loss of leaf litter were assessed. The field study was performed on an abandoned rubber plantation near Manaus (Amazonia, Brazil). The measurement endpoints were abundance of the soil fauna and mass loss of leaf litter. In the TMEs carbendazim caused a decrease in the abundance of the introduced earthworm Pontoscolex corethrurus and, partly, of the milliped Trigoniulus corallinus. In the field carbendazim decreased the abundance of the native earthworm Andiorrhinus amazonius. Lambda-cyhalothrin was toxic to isopods and millipedes in the TMEs, whereas no effect on arthropods was detected in the field. Organic matter breakdown measured as mass loss of leaf litter in TMEs over time revealed that the two tested agrochemicals can have an impact on decomposition at field relevant concentrations. In the field the results were less obvious, due to spatial heterogeneity through which possible effects of the tested agrochemicals could have been masked.     

Community structure of oribatid mites in relation to elevation and geology on the slope of Mount Kinabalu,Sabah, Malaysia

The patterns of oribatid communities were investigated in relation to elevations (700, 1700, 2700 and 3100 m a.s.l.) and geological substrates (i.e. non-ultrabasic and ultrabasic rocks) on the slopes of Mt. Kinabalu, Sabah, Malaysia. The density and morphospecies richness of oribatid mites were greater in the non-ultrabasic plot than in the ultrabasic plot at each of the same elevations. The density and richness of Oribatid mites decreased with elevation on both substrates, but the effects of elevation on the density on non-ultrabasic were less significant than on the ultrabasic substrate. Oribatid mite density correlated positively with the concentration of soil organic phosphorus and negatively with that of exchangeable Ca in soil. The richness of morphospecies of oribatid mites positively correlated with phosphorus concentration in litter, above-ground biomass, tree diversity and litterfall amount, and negatively correlated with elevation and Ca in soil. Morphospecies from families Galumnidae, Otocepheidae, Haplozetidae and Scheloribatidae were dominant in each plot. Canonical correspondence analysis (CCA) showed the importance of elevation for the community structure of oribatid mite. In conclusion, total density or morphospecies richness of oribatid communities was influenced by both geology and elevation, and morphospecies composition was strongly influenced by elevation.     

Decomposition and nitrogen dynamics of litter in peat soils from two climatic regions under different temperature regimes

Soil temperature is a major factor affecting organic matter decomposition and thus, global warming may accelerate decomposition processes. However, it remains unclear whether the effects will be similar in climatically different regions. The effects of soil temperatures of 5, 10 and 15 °C on the decomposition of Scots pine (Pinus sylvestris L.) needles were assessed in a 1-year (360 days) growth chamber experiment. Intact peat cores from two climatically different peatland sites (southern and northern Finland) were used as the incubation environments. Needles were incubated in litter bags beneath the living moss layer, and mass loss and nitrogen (N) concentration were determined at 60-day intervals. The rate of mass loss from the needles over time was clearly lower in the 5 °C treatment than at the higher temperatures. Mass loss was strongly related to the accumulated soil temperature sum. In temperatures higher than 5 °C, mass losses were higher in the northern peat. Also, the limit value of decomposition (asymptotic maximum mass loss) was slightly higher in the northern peat (92%), than in the southern peat (87%). The N concentration increased up to a mass loss of 50–60%, whereupon it decreased, while the amount of N (as a percentage of the original amount) remained unchanged until a mass loss of 50–60%, whereupon it decreased linearly. It seems that increasing soil temperatures may result in slightly higher rates of needle litter mass loss and consequent N release in northern peat than in southern peat. The faster decomposition in higher temperatures in the northern peat, together with the slightly higher maximum mass loss value, imply that with climatic warming, susceptibility of boreal peatlands for becoming sources of carbon to the atmosphere may increase towards north.     

C and N mineralization and microbial biomass in heavy-metal contaminated soil

Heavy metals such as arsenic (As), lead (Pb), copper (Cu) and zinc (Zn) can be found in large concentrations in mine spills in Mexico. Interest in contamination by these heavy metals has increased recently as they can change the functioning of soil ecosystems qualitatively and quantitatively. They disturb the activities of soil fauna and contaminate drinking water in large parts of the world, which severely affects human health. Little, however, is known how heavy metals might affect the biological functioning of a soil. Soil was sampled from eight locations along a gradient of heavy-metal contamination with distance from a mine in San Luis Potosí (Mexico) active since about 1800 AD. Microbial biomass was determined with the original chloroform fumigation incubation (CFI) as well as extraction (CFE) techniques and the substrate induced respiration (SIR) technique while C and N mineralization were measured. Total concentrations of As in the top 0–10 cm soil layer ranged from 8 to 22992 mg kg^–1, from 31 to 1845 mg kg^–1 for Pb, from 27 to 1620 mg kg^–1 for Cu and from 81 to 4218 mg kg^–1 for Zn. There was a significant negative correlation (P < 0.0001) between microbial biomass, soil organic carbon, total N and C mineralization and the heavy metal content of the soil. The microbial biomass C to organic C ratio, which varied from 0.4 to 1.9%, specific respiratory activity (qCO₂), and oxidation of NO₂^– were not affected by heavy metals. It was found that long-term contamination of soil with heavy metals had an adverse effect on the amount of soil microorganisms as evidenced by a marked decrease in microbial biomass C, but not some of their characteristics. According to principal components analysis (PCA), the correlation matrix showed three distinct factors explaining 71% of the variance. A first factor including heavy metals (As, Pb, Cu and Zn) with a negative loading and total N, organic C, soil microbial biomass with a positive loading characterized the soil organic matter and contamination status. Loam and sand combined for the second factor characterizing the textural classification while the third factor was loaded by CEC and clay content.     

Patch-dynamics and population structure of nematodes and tardigrades on Antarctic nunataks

Assemblages of hydrophilic microfauna (nematodes, rotifers and tardigrades) in moss-cushions from four ice-free mountain peaks (nunataks) in East Antarctica were studied. Nematodes occurred in 64%, tardigrades in 32% and rotifers in 82% of 91 samples taken in moss. No animals were found in 8% of the samples. The differences in occurrence of animals among cushions are probably due to stochastic colonization processes. The structure of nematode populations indicates that they exhibit different developmental stages in the various samples. Competition or predation appears to influence the abundances of coexisting populations. Predation by tardigrades on rotifers might occur, while the nematodes appear to be competitors rather than preys of the tardigrades.     

and Its Expression in Pichia pastoris

利用RT-PCR技术扩增出草鱼(Ctenopharyngodon idellus)的leptin基因，将leptin基因克隆至真核表达载体pPIC9K，电穿孔转化GS115菌株, 经G418筛选和甲醇诱导后，对表达产物进行SDS-PAGE琼脂糖凝胶电泳和Western blot分析。结果表明，草鱼leptin基因cDNA序列由438个核苷酸组成，编码146个氨基酸组成的多肽（GenBank登陆号AY551335），与鲤鱼(Cyprinus carpio)leptin基因相比， 核苷酸和氨基酸的同源性为99%； 与人、猪和鼠相比，核苷酸同源性分别为84%、86%和95%, 氯基酸的同源性分别为84%、82%和96%；与河豚(Takifugu rubripes)相比，氨基酸具有较大的差异，仅有9％的同源性，表明leptin在物种的进化上具有一定的差异；实现了草鱼leptin基因在毕赤酵母(Pichia pastoris)中的表达，表达蛋白的分子量约为16 kD， Western blot 分析表明，表达产物具有一定的免疫学活性。     

in Gastrointestinal Tracts of Chicken

以30 d Arbor Acre (AA)肉鸡为模型，采集胃肠道样品，采用相对定量RT-PCR方法研究了生长抑素(SS ) mRNA在鸡胃肠道分布的组织特异性。结果表明，腺胃SS mRNA的表达量高于所有肠段，差异极显著 (P＜0.01)；十二指肠和空肠SS mRNA的表达丰度相似，二者均显著高于回肠(P＜0.05)。定性研究显示，肌胃和结直肠均有SS mRNA表达，但其表达量显著低于腺胃、十二指肠、空肠和回肠。SS mRNA在主要的消化(腺胃)、吸收(十二指肠和空肠前段)器官的高丰度表达，提示SS对营养物质的消化吸收具有重要的调控作用。