Biotic contexts alter metal sequestration and AMF effects on plant growth in soils polluted with heavy metals

Investigating how arbuscular mycorrhizal fungi (AMF)-plant interactions vary with edaphic conditions provides an opportunity to test the context-dependency of interspecific interactions. The relationship between AMF and their host plants in the context of other soil microbes was studied along a gradient of heavy metal contamination originating at the site of zinc smelters that operated for a century. The site is currently under restoration. Native C3 grasses have reestablished, and C4 grasses native to the region but not the site were introduced. Interactions involving the native mycorrhizal fungi, non-mycorrhizal soil microbes, soil, one C3 grass (Deschampsia flexuosa), and one C4 grass (Sorghastrum nutans) were investigated using soils from the two extremes of the contamination gradient in a full factorial greenhouse experiment. After 12 weeks, plant biomass and root colonization by AMF and non-mycorrhizal microbes were measured. Plants from both species grew much larger in soil from low-contaminated (LC) origin than high-contaminated (HC) origin. For S. nutans, the addition of a non-AMF soil microbial wash of either origin increased the efficacy of AMF from LC soils but decreased the efficacy of AMF from HC soils in promoting plant growth. Furthermore, there was high mortality of S. nutans in HC soil, where plants with AMF from HC died sooner. For D. flexuosa, plant biomass did not vary with AMF source or the microbial wash treatment or their interaction. While AMF origin did not affect root colonization of D. flexuosa by AMF, the presence and origin of AMF did affect the number of non-mycorrhizal (NMF) morphotypes and NMF root colonization. Adding non-AMF soil biota reduced Zn concentrations in shoots of D. flexuosa. Thus the non-AMF biotic context affected heavy metal sequestration and associated NMF in D. flexuosa, and it interacted with AMF to affect plant biomass in S. nutans. Our results should be useful for improving our basic ecological understanding of the context-dependency of plant-soil interactions and are potentially important in restoration of heavy-metal-contaminated sites.     

Specificity between Neotropical tree seedlings and their fungal mutualists leads to plant-soil feedback

A growing body of evidence obtained largely from temperate grassland studies suggests that feedbacks occurring between plants and their associated soil biota are important to plant community assemblage. However, few studies have examined the importance of soil organisms in driving plant-soil feedbacks in forested systems. In a tropical forest in central Panama, we examined whether interactions between tree seedlings and their associated arbuscular mycorrhizal fungi (AMF) lead to plant-soil feedback. Specifically, do tropical seedlings modify their own AMF communities in a manner that either favors or inhibits the next cohort of conspecific seedlings (i.e., positive or negative feedback, respectively)? Seedlings of two shade-tolerant tree species (Eugenia nesiotica, Virola surinamensis) and two pioneer tree species (Luehea seemannii, Apeiba aspera) were grown in pots containing identical AMF communities composed of equal amounts of inoculum of six co-occurring AMF species. The different AMF-host combinations were all exposed to two light levels. Under low light (2% PAR), only two of the six AMF species sporulated, and we found that host identity did not influence composition of AMF spore communities. However, relative abundances of three of the four AMF species that produced spores were influenced by host identity when grown under high light (20% PAR). Furthermore, spores of one of the AMF species, Glomus geosporum, were common in soils of Luehea and Eugenia but absent in soils of Apeiba and Virola. We then conducted a reciprocal experiment to test whether AMF communities previously modified by Luehea and Apeiba differentially affected the growth of conspecific and heterospecific seedlings. Luehea seedling growth did not differ between soils containing AMF communities modified by Luehea and Apeiba. However, Apeiba seedlings were significantly larger when grown with Apeiba-modified AMF communities, as compared to Apeiba seedlings grown with Luehea-modifed AMF communities. Our experiments suggest that interactions between tropical trees and their associated AMF are species-specific and that these interactions may shape both tree and AMF communities through plant-soil feedback.     

Soil feedback of exotic savanna grass relates to pathogen absence and mycorrhizal selectivity

Enemy release of exotic plants from soil pathogens has been tested by examining plant-soil feedback effects in repetitive growth cycles. However, positive soil feedback may also be due to enhanced benefit from the local arbuscular mycorrhizal fungi (AMF). Few studies actually have tested pathogen effects, and none of them did so in arid savannas. In the Kalahari savanna in Botswana, we compared the soil feedback of the exotic grass Cenchrus biflorus with that of two dominant native grasses, Eragrostis lehmanniana and Aristida meridionalis. The exotic grass had neutral to positive soil feedback, whereas both native grasses showed neutral to negative feedback effects. Isolation and testing of root-inhabiting fungi of E. lehmanniana yielded two host-specific pathogens that did not influence the exotic C. biflorus or the other native grass, A. meridionalis. None of the grasses was affected by the fungi that were isolated from the roots of the exotic C. biflorus. We isolated and compared the AMF community of the native and exotic grasses by polymerase chain reaction-denaturing gradient gel elecrophoresis (PCR-DGGE), targeting AMF 18S rRNA. We used roots from monospecific field stands and from plants grown in pots with mixtures of soils from the monospecific field stands. Three-quarters of the root samples of the exotic grass had two nearly identical sequences, showing 99% similarity with Glomus versiforme. The two native grasses were also associated with distinct bands, but each of these bands occurred in only a fraction of the root samples. The native grasses contained a higher diversity of AMF bands than the exotic grass. Canonical correspondence analyses of the AMF band patterns revealed almost as much difference between the native and exotic grasses as between the native grasses. In conclusion, our results support the hypothesis that release from soil-borne enemies may facilitate local abundance of exotic plants, and we provide the first evidence that these processes may occur in arid savanna ecosystems. Pathogenicity tests implicated the involvement of soil pathogens in the soil feedback responses, and further studies should reveal the functional consequences of the observed high infection with a low diversity of AMF in the roots of exotic plants.     

Soil biota drive expression of genetic variation and development of population-specific feedbacks in an invasive plant

Invasive plant species alter soils in ways that may affect the success of subsequent generations, creating plant-soil feedbacks. Ailanthus altissima is an invasive tree introduced two centuries ago to North America. We hypothesized that geographically distinct populations of A. altissima have established feedbacks specific to their local environment, due to soil communities cultivated by A. altissima. We collected seeds and soils from three populations in the eastern United States, and in the greenhouse reciprocally planted all families in all collected soils as well as in a control mixed soil, and in soils that had been irradiated for sterilization. There were positive plant-soil feedbacks for two populations in the live field-collected soils, but strong negative feedbacks for the third population. There were no population-level performance differences or feedbacks in the sterilized population locale soils, supporting a soil biotic basis for feedbacks and for the expression of genetic differentiation in A. altissima. If populations of Ailanthus altissima vary in the extent to which they benefit from and promote these plant-soil biota feedbacks, the interaction between invader and invaded community may be more important in determining the course of invasion than are the characteristics of either alone.     

Mycorrhizal fungal identity and diversity relaxes plant-plant competition

There is a great interest in ecology in understanding the role of soil microbial diversity for plant productivity and coexistence. Recent research has shown increases in species richness of mutualistic soil fungi, the arbuscular mycorrhizal fungi (AMF), to be related to increases in aboveground productivity of plant communities. However, the impact of AMF richness on plant-plant interactions has not been determined. Moreover, it is unknown whether species-rich AMF communities can act as insurance to maintain productivity in a fluctuating environment (e.g., upon changing soil conditions). We tested the impact of four different AMF taxa and of AMF diversity (no AMF, single AMF taxa, and all four together) on competitive interactions between the legume Trifolium pratense and the grass Lolium multiflorum grown under two different soil conditions of low and high sand content. We hypothesized that more diverse mutualistic interactions (e.g., when four AMF taxa are present) can ease competitive effects between plants, increase plant growth, and maintain plant productivity across different soil environments. We used quantitative PCR to verify that AMF taxa inoculated at the beginning of the experiment were still present at the end. The presence of AMF reduced the competitive inequality between the two plant species by reducing the growth suppression of the legume by the grass. High AMF richness enhanced the combined biomass production of the two plant species and the yield of the legume, particularly in the more productive soil with low sand content. In the less productive (high sand content) soil, the single most effective AMF had an equally beneficial effect on plant productivity as the mixture of four AMF. Since contributions of single AMF to plant productivity varied between both soils, higher AMF richness would be required to maintain plant productivity in heterogeneous environments. Overall this work shows that AMF diversity promotes plant productivity and that AMF diversity can act as insurance to sustain plant productivity under changing environmental conditions.     

丛枝菌根对土壤-植物系统中重金属迁移转化的影响

丛枝菌根真菌(arbuscular mycorrhizal fungi, AMF)是一类在自然和农业生态系统中广泛存在并能与多数陆生植物形成共生关系的土壤真菌,在重金属污染土壤中对宿主植物的生长及吸收累积重金属具有重要影响,因而对污染土壤的生物修复具有潜在应用价值。以重金属从根际土壤进入植物并在植物体内再分配过程为主线,介绍丛枝菌根在这一过程中对重金属环境行为,特别是根际土壤中重金属赋存形态及植物吸收重金属的影响。最后,对丛枝菌根影响植物重金属耐性机制研究前沿和菌根修复技术的应用前景进行展望。     

Volatile chemicals from leaf litter are associated with invasiveness of a neotropical weed in Asia

Some invasive plant species appear to strongly suppress neighbors in their nonnative ranges but much less so in their native range. We found that in the field in its native range in Mexico, the presence of Ageratina adenophora, an aggressive Neotropical invader, was correlated with higher plant species richness than found in surrounding plant communities where this species was absent, suggesting facilitation. However, in two nonnative ranges, China and India, A. adenophora canopies were correlated with much lower species richness than the surrounding communities, suggesting inhibition. Volatile organic compound (VOC) signals may contribute to this striking biogeographical difference and the invasive success of A. adenophora. In controlled experiments volatiles from A. adenophora litter caused higher mortality of species native to India and China, but not of species native to Mexico. The effects of A. adenophora VOCs on seedling germination and growth did not differ between species from the native range and species from the nonnative ranges of the invader. Litter from A. adenophora plants from nonnative populations also produced VOCs that differed quantitatively in the concentrations of some chemicals than litter from native populations, but there were no chemicals unique to one region. Biogeographic differences in the concentrations of some volatile compounds between ranges suggest that A. adenophora may be experiencing selection on biochemical composition in its nonnative ranges.     

Influence of invasive plant Coreopsis grandiflora on functional diversity of soil microbial communities

Biological invasions are increasingly attracting the ecologists' attention. Invasive plants threaten the natural ecosystems not only by competing with the native plants, but also by altering the structure and function of soil microbial communities belowground. In this study, we studied the effects of the invasive plant Coreopsis grandiflora (C. grandiflora) on the functional diversity of soil microbial communities in Laoshan mountain in the province of Shandong, North of China. We sampled soil from plots that were invaded or not invaded by C. grandiflora. The functional diversity of microbial communities in the sampled soils was assessed by the Biolog procedure test. By the ANOVA analysis of average well color development (AWCD), Shannon index (H'), Shannon evenness (E), principle components analysis of the level physiological profiles (CLPP) and correlation analysis between the studied parameters, we found that the invasive species C. grandiflora enhanced the functional diversity of soil microbial communities where the habitat was invaded by the C. grandiflora. The study indicated thatthe successful invasive plants have profound effects on the function of soil microbial communities.     

Novel weapons: invasive plant suppresses fungal mutualists in America but not in its native Europe

Why some invasive plant species transmogrify from weak competitors at home to strong competitors abroad remains one of the most elusive questions in ecology. Some evidence suggests that disproportionately high densities of some invaders are due to the release of biochemicals that are novel, and therefore harmful, to naive organisms in their new range. So far, such evidence has been restricted to the direct phytotoxic effects of plants on other plants. Here we found that one of North America's most aggressive invaders of undisturbed forest understories, Alliaria petiolata (garlic mustard) and a plant that inhibits mycorrhizal fungal mutualists of North American native plants, has far stronger inhibitory effects on mycorrhizas in invaded North American soils than on mycorrhizas in European soils where A. petiolata is native. This antifungal effect appears to be due to specific flavonoid fractions in A. petiolata extracts. Furthermore, we found that suppression of North American mycorrhizal fungi by A. petiolata corresponds with severe inhibition of North American plant species that rely on these fungi, whereas congeneric European plants are weakly affected. These results indicate that phytochemicals, benign to resistant mycorrhizal symbionts in the home range, may be lethal to na?ve native mutualists in the introduced range and indirectly suppress the plants that rely on them.     

Variation in resource availability changes the impact of invasive thistles on native bunchgrasses.

The threat posed by invasive nonnative plants to native plant populations is one of the largest challenges facing both conservation biology and restoration ecology. California has been highly impacted by invaders, although many relict stands of native plants are found on shallow, rocky soils with limited resources. The abiotic conditions of these sites may strongly influence the performance of an invasive plant and its effect on resident native species. In addition, the maturity of native plants in these sites may modulate an invader's impact; larger, well-established plants may be better able to resist invaders. In this study we examined how the impact of an invasive thistle (Centaurea solstitialis) on a native perennial bunchgrass (Nassella pulchra) changed in response to variation in soil depth, soil water availability, and bunchgrass maturity. We measured plant performance in terms of survival, growth, reproduction, and predawn water potential. We found that soil depth, water availability, and bunchgrass maturity acted in concert to influence the impact of the invasive thistle on the native bunchgrass. Both species performed better in deep soils, especially during dry years. The combination of shallow soil and low water availability reduced C. solstitialis performance and ameliorated its negative effect on N. pulchra growth and reproduction. Higher water availability resulted in a stronger negative effect of C. solstitialis on N. pulchra in both shallow and deep soils. However, as N. pulchra matured and increased in size, we saw a steady decline in C. solstitialis growth and reproductive output. Higher water availability increased the performance of C. solstitialis in shallow soils. C. solstitialis may thus have a stronger impact on N. pulchra and be more able to invade relict stands of N. pulchra in shallow soils during high-rainfall years. However, established stands of N. pulchra appear to be more resistant to invasion by C. solstitialis as N. pulchra plants grow older and larger.     

Escape from competition: neighbors reduce Centaurea stoebe performance at home but not away

The greater abundance of some exotic plants in their nonnative ranges might be explained in part by biogeographic differences in the strength of competition, but these competitive effects have not been experimentally examined in the field. We compared the effects of neighbors on the growth and reproduction of spotted knapweed (Centaurea stoebe) in Europe, where it is native, and in Montana, where it is invasive. There were strong negative competitive effects of neighboring vegetation on C. stoebe growth and reproduction in Europe. In contrast, identical experiments in Montana resulted in insignificant impacts on C. stoebe. Although the mechanisms that produce this dramatic biogeographic difference in competitive outcome remain unknown, our results indicate that differences in net competitive interactions between ranges may contribute to the striking dominance of C. stoebe in parts of North America.     

云南会泽废弃铅锌矿区植物丛枝菌根和深色有隔内生真菌研究

用碱解离、酸性品红染色法对云南省会泽县者海镇废弃铅锌矿区的17科21种植物的丛枝菌根状况进行了调查,结果发现,15种植物形成典型的丛枝菌根,占所调查植物的71%;2种植物不确定是否形成丛枝菌根,占所调查植物的10%;4种植物没有形成丛枝菌根,占所调查植物的19%.用湿筛沉淀法从这些植物根际土壤中共分离鉴定出了4属20种丛枝菌根真菌(AMF),即无梗囊霉属(Acaulospora)4种,球囊霉属(Glomus)14种,巨孢囊霉属(Gigaspo-ra)1种,盾巨孢囊霉属(Scutellospora)1种;其中,球囊霉属分离频率为77%,是样地的优势属.在AMF中,疣突球囊霉(G.verruculosum)分离频率最高,在20种植物的根际土中都有发现;此外,聚生球囊霉(G.fasciculatum)的相对多度最大,为56%,具有最强的产孢能力.同时,在13种植物的根中发现了深色有隔内生真菌(DSE),占调查植物的62%,其中,10种植物同时被DSE和AMF感染.本调查研究表明,AMF和DSE能普遍存在于Pb、Zn重金属污染土壤中.     

2种入侵植物对根际土壤微生物种群及代谢的影响

选取2种入侵植物（一年蓬Annual Fleabane及加拿大蓬Erigeron Canadensis）及本土植物（艾蒿Artemisia argyi）的根际土壤微生物种群为研究对象,以分析不同根际土壤微生物种群的数量及测定根际土壤微生物酶活的活性为切入点来探析入侵植物对土壤微生物的影响及其响应机制。结果显示：2种入侵植物显著增加了其根际土壤中的细菌的数量,而显著抑制了真菌与放线菌的数量。另外,入侵植物一年蓬显著抑制其根际土壤中纤维素酶的活性,而3种植物的根际土壤硝酸还原酶活性无显著差异。入侵植物对其他3种土壤酶（即转化酶、脲酶及酸性磷酸酶）活性的影响却呈现出截然相反的影响,即一年蓬显著增加了3种根际土壤酶的活性,而加拿大蓬却显著减少了3种根际土壤酶的活性。导致这种现象的原因可能是不同入侵植物的根系释放不同的化学物质进而对土壤微生物的数量和活性造成不同的影响。     

喀斯特生境下AMF侵染对任豆生长的影响

喀斯特地区土壤退化,植被定植更新困难,丛枝菌根真菌(Arbuscular mycorrhizal fungi,AMF)具有增强植物养分吸收能力和抵抗逆境胁迫能力。研究喀斯特生境下植物与AMF共生效果,选择优势菌种促进喀斯特植被恢复,对于提高植物定植成活率具有重要作用。以豆科植物任豆(Zenia insignis)幼苗为试验材料,盆栽条件下,选取喀斯特优势菌种-摩西球囊霉(Funneliformis mosseae)、根内球囊霉(Rhizophagus intraradices),2种菌根真菌混合菌剂进行接种,培养180 d,研究贫瘠喀斯特土壤生境和养分较高的滇柏林下土壤生境下AMF对任豆生长影响。结果表明:摩西球囊霉、根内球囊霉和混合接种均能侵染任豆根系,幼嫩根系更易侵染,木质化根系侵染率下降。接种摩西球囊霉,贫瘠喀斯特土壤生境下,株高、地径、地上生物量、地下生物量和总生物量分别提高68.92%、56.18%、83.90%、42.20%和67.34%;养分较高的滇柏林下喀斯特土壤生境下,株高、地上生物量、地下生物量和总生物量分别提高48.05%、6.77%、7.92%和8.89%;根内球囊霉处理接种效应低于摩西球囊霉和混合接种处理,对生物量增长为负效应,混合接种处理接种效应介于单接种之间,摩西球囊霉接种效果优于根内球囊霉和混合接种。摩西球囊霉在贫瘠喀斯特土壤生境下发挥的促生效应优于养分较高的喀斯特土壤,可作为喀斯特侵蚀区植被恢复菌根真菌干扰途径的优势菌种,混合接种作为接种剂具有单接种兼容效应。     

Ableitung ökotoxikologisch begründeter Bodenqualitätskriterien am Beispiel Arsen

Background

A vast amount of data on the concentration of substances in soils is being generated in environmental monitoring programmes. This data has to be assessed in terms of effects on terrestrial biota. Benchmarks for such an assessment have been missing up to now. This gap has been filled by the development of a concept for the derivation of ecotoxicologically based soil quality criteria (SQC) and the application of this concept to environmentally relevant substances.

Objectives

The derivation method was applied to arsenic. The derivation process was described in detail from the compilation of data regarding effects on microbial processes, plants, and soil invertebrates to the formulation of a recommendation for an SQC.

Methods

The conceptual framework for the derivation of ecotoxicologically based SQC was developed after reviewing national and international concepts for the assessment of the effects of substances on terrestrial communities. For the assessment of arsenic effects, all relevant biotic processes and trophic levels have been taken into account. Moreover, soil characteristics that may affect the toxicity to biota have been reviewed.

Results

Effects of arsenic on biota may occur at soil background concentrations or slightly above. There are only a few studies in which the influence of soil characteristics, such as soil pH, soil texture, and soil organic carbon, on the effects of arsenic has been investigated systematically. That is why the SQC was oriented on soil background concentrations for sandy soils (5 mg As/kg soil dw). A comparison of the SQC derived in this work with ecotoxicologically based soilquality values from Canada, Denmark, the Netherlands, and the United States shows an average value of 16 mg As/kg, with a range between 5 and 34 mg/kg. The variations in the soil quality values are mainly due to differences in the derivation approaches.

Conclusions and Outlook

The concept applied is considered suitable for the derivation of SQC designed to protect terrestrial communities against direct effects of substances in soils. Therefore, its use is planned to derive SQC for other environmentally relevant substances. In addition to compensation factors, statistical methods should be taken into account to compensate for risks which cannot be quantified. Furthermore, there is the need for an assessment of indirect effects of soil contaminants on birds and mammals.     

Cars, Cows, and Checkerspot Butterflies: Nitrogen Deposition and Management of Nutrient-Poor Grasslands for a Threatened Species

Abstract: Nutrient-poor, serpentinitic soils in the San Francisco Bay area sustain a native grassland that supports many rare species, including the Bay checkerspot butterfly ( Euphydryas editha bayensis ). Nitrogen (N) deposition from air pollution threatens biodiversity in these grasslands because N is the primary limiting nutrient for plant growth on serpentinitic soils. I investigated the role of N deposition through surveys of butterfly and plant populations across different grazing regimes, by literature review, and with estimates of N deposition in the region. Several populations of the butterfly in south San Jose crashed following the cessation of cattle grazing. Nearby populations under continued grazing did not suffer similar declines. The immediate cause of the population crashes was rapid invasion by introduced annual grasses that crowded out the larval host plants of the butterfly. Ungrazed serpentinitic grasslands on the San Francisco Peninsula have largely resisted grass invasions for nearly four decades. Several lines of evidence indicate that dry N deposition from smog is responsible for the observed grass invasion. Fertilization experiments have shown that soil N limits grass invasion in serpentinitic soils. Estimated N deposition rates in south San Jose grasslands are 10–15 kg  N/ha/year; Peninsula sites have lower deposition, 4–6 kg N/ha/year. Grazing cattle select grasses over forbs, and grazing leads to a net export of N as cattle are removed for slaughter. Although poorly managed cattle grazing can significantly disrupt native ecosystems, in this case moderate, well-managed grazing is essential for maintaining native biodiversity in the face of invasive species and exogenous inputs of N from nearby urban areas.     

Bioavailability,mobility, and toxicity of Cu in soils around the Dexing Cu mine in China

Total concentration is not a reliable indicator of Cu mobility or its bioavailability in soils. The chemical fraction determines the behavior of Cu in soils and hence its toxicity for terrestrial biota. We used the sequential extraction procedure and barley toxicity tests to examine the transfer of Cu in soils around the Dexing Cu mine and to make an ecological risk assessment of Cu in this area. The bioavailable Cu (exchangeable Cu and carbonate-bound Cu) in each soil profile did not change significantly with soil depth, indicating that the Cu itself was vertically mobile and thus potentially a higher risk to the environment. Cu toxicity and bioaccumulation in plants varied with the soil physicochemical characteristics [e.g., pH, clay content, and cation exchange capacity (CEC)] and the level of bioavailable Cu. Multiple regression analysis revealed that bioavailable Cu and CEC could be used to predict Cu toxicity to barley and that other characteristics (such as soil pH, clay content, or total organic carbon) did not predict the risk of toxicity as well as CEC. For the soil to be suitable for agriculture use, treatment of the local soil contamination with guest soil reclamation and phytoremediation with heavy metal-resistant plants would be necessary.     

外来入侵植物化感作用与土壤相互关系研究进展

文章以全球普遍关注的外来植物入侵为背景,综述了外来入侵植物化感物质与土壤化学性质(土壤pH,土壤有机碳和有机质,土壤中的化学元素等)、土壤生物群落(土壤微生物、土壤动物)的关系。并探讨了未来研究需要加强的几个方面,包括将化感作用与野外实际情况相结合研究,更真实的反映田间状况;进一步探讨微生物在化感物质生物变化过程中的作用及其机理;化感物质作为信号物质如何影响微生物代谢,从而影响地上植物;深入研究化感作用与土壤之间的相互关系,为入侵植物生物防治的安全性提供理论指导等方面。     

酸雨对外来植物入侵的影响

酸雨和外来种入侵都是全球关注的问题。结合外来入侵植物的生态适应特性以及酸雨的危害特征,系统分析了酸雨对外来植物入侵产生的影响。酸雨对外来植物入侵的影响是复杂多样的。酸雨导致群落冠层稀疏,群落透光率增加,加之氮沉降后土壤、水体氮素的增加,有利于生长力强的外来喜阳植物入侵;酸雨加速土壤酸化,促使基本离子淋失以及A1毒等危害植物的生长发育,植物的内源激素以及化感作用发生改变,适应力和耐受力强的外来植物在与本地植物竞争中处于相对优势而成为入侵种;酸雨以及外来植物入侵改变了土壤微生物群落结构,影响本地植物的生长而促使外来植物的入侵。