Terrestrial Protozoa*

SYNOPSIS Litters and soils are special interstitial ecosystems containing water in surface films and pore spaces, large amounts of organic matter, and are subject to extreme moisture and temperature fluctuations. “Terrestrial protozoa” are ubiquitous limnetic species tolerant of high CO₂ tensions and possessing efficient encystment mechanisms. Protozoa exploit tiny microhabitats unavailable to larger animals (e.g. nematodes). Naked amebae dominate due to their flexible bodies and interface locomotion. Small flagellates may be abundant, especially in litters. Ciliates are less numerous but their species composition indicates the degree of moisture of the habitat, as do the slower-growing testacea which become prominent in regions of slow decomposition (conifer and tundra biomes). Protozoa promote decomposition by enhancing bacterial metabolism, eating excess bacteria, and excreting simple compounds returnable to plants. Large populations, especially amebae, exploit the abundant bacterial flora of plant root zones (rhizosphere). One protozoon, Colpoda cucullus, has successfully invaded the surfaces of vegetation.     

Protozoa,Nematoda and Lumbricidae in the rhizosphere of Hordelymus europeaus (Poaceae): faunal interactions,response of microorganisms and effects on plant growth

Interactions among protozoa (mixed cultures of ciliates, flagellates and naked amoebae), bacteria-feeding nematodes (Pellioditis pellio Schneider) and the endogeic earthworm species Aporrectodea caliginosa (Savigny) were investigated in experimental chambers with soil from a beechwood (Fagus sylvatica L.) on limestone. Experimental chambers were planted with the grass Hordelymus europeaus L. (Poaceae) and three compartments separated by 45-m mesh were established: rhizosphere, intermediate and non-rhizosphere. The experiment lasted for 16 weeks and the following parameters were measured at the end of the experiment: shoot and root mass of H. europaeus, carbon and nitrogen content in shoots and roots, density of ciliates, amoebae, flagellates and nematodes, microbial biomass (SIR), basal respiration, streptomycin sensitive respiration, ammonium and nitrate contents, phosphate content of soil compartments. In addition, leaching of nutrients (nitrogen and phosphorus) and leachate pH were measured at regular intervals in leachate obtained from suction cups in the experimental chambers. Protozoa stimulated the recovery of nitrifying bacteria following defaunation (by chloroform fumigation) and increased nitrogen losses as nitrate in leachate. In contrast, protozoa and nematodes reduced leaching of phosphate, an effect ascribed to stimulation of microbial growth early in the experiment. Earthworms strongly increased the amount of extractable mineral nitrogen whereas it was strongly reduced by protozoa and nematodes. Both protozoa and nematodes reduced the stimulatory effect of earthworms on nitrogen mineralization. Microbial biomass, basal respiration, and numbers of protozoa and nematodes increased in the vicinity of the root. Protozoa generally caused a decrease in microbial biomass whereas nematodes and earthworms reduced microbial biomass only in the absence of protozoa. None of the animals studied significantly affected basal respiration, but specific respiration of microorganisms (O₂ consumption per unit biomass) was generally higher in animal treatments. The stimulatory effect of nematodes and earthworms, however, occurred only in the absence of protozoa. The sensitivity of respiration to streptomycin suggested that protozoa selectively grazed on bacterial biomass but the bacterial/fungal ratio appeared to be unaffected by grazing of P. pellio. Earthworms reduced root biomass of H. europaeus, although shoot biomass remained unaffected, and concentrations of nitrogen in shoots and particularly in roots were strongly increased by earthworms. Both nematodes and protozoa increased plant biomass, particularly that of roots. This increase in plant biomass was accompanied by a marked decrease in nitrogen concentrations in roots and to a lesser extent in shoots. Generally, the effects of protozoa on plant growth considerably exceeded those of nematodes. It is concluded that nematodes and protozoa stimulated plant growth by non-nutritional effects, whereas the effects of earthworms were caused by an increase in nutrient supply to H. europaeus.     

Protozoan Grazing Increases Mineralization of Naphthalene in Marine Sediment

Bacterial decomposition of organic matter is frequently enhanced when protozoa are present. Various mechanisms have been proposed to account for this phenomenon, including effects associated with grazing by protozoa (such as increased recycling of limiting nutrients, removal of senescent cells, or reduction of competition among bacteria) and indirect effects of grazers (such as excretion of bacterial growth factors). Few studies have examined the role of protozoa in bacterial degradation of xenobiotic compounds in sediment containing a natural community of microbes. The effect of protozoa on mineralization of naphthalene was investigated in this study. Laboratory experiments were conducted using field-contaminated estuarine sediment, with the indigenous microbial populations. Mineralization of naphthalene was up to four times greater in treatments with actively grazing protozoa than in treatments containing the grazing inhibitor cytochalasin B. Control experiments confirmed that the grazing inhibitor was not toxic to ciliates but did prevent them from grazing. The grazing inhibitor did not affect growth rates of a mixed culture of sediment bacteria or a pure polycyclic-aromatic-hydrocarbon-degrading strain. Once grazing had been inhibited, supplementing treatments with inorganic N and P, glucose, or additional protozoa failed to stimulate naphthalene mineralization. Naphthalene-degrading bacteria were four to nine times less abundant when protozoan grazing was suppressed. We suggest that protozoa enhance naphthalene mineralization by selectively grazing on those sediment bacteria that ordinarily would outcompete naphthalene-degrading bacteria.     

Simultaneous consumption of bacteria and dissolved organic matter byTetrahymena pyriformis

The addition of ciliated protozoa to aquatic microcosms and bench-scale sewage treatment plants increases decomposition rates. This is surprising, inasmuch as protozoa consume bacteria, which are the primary decomposers. One possible mechanism of the increase in decomposition rate is the direct consumption of dissolved organic matter by protozoa that are feeding primarily on bacteria. This possibility was explored experimentally in two-stage continuous cultures, with glucose limitingEscherichia coli in the first stage andE. coli limitingTetrahymena pyriformis in the second. Glycine and histidine were the test nutrients. The results of adding them to the second stages suggested that direct uptake by ciliates does not affect the dynamics of dissolved amino acids in pelagic environments or activated sludge plants. Ciliates might, however, affect the dynamics of amino acid pools in environments high in nutrients and ciliates, perhaps including some microenvironments near decomposing material or in benthic sediments. Direct uptake of dissolved amino acids by ciliates probably does not affect ciliate or bacterial populations substantially.     

Simulated herbivory effects on rhizosphere organisms in pea (Pisum sativum) depended on phosphate

The effects of simulated aboveground herbivory and phosphate addition to soil on rhizosphere organisms (arbuscular mychorrhiza (AM), Rhizobium spp., bacteria, protozoa and nematodes) were studied in a 2 by 2 factorial designed pot experiment with Pea plants (Pisum sativum). Measurements were performed on 24 day old plants that were still in the nutrient acquisition phase before flowering. AM colonization and bacterial feeding nematodes were stimulated by high simulated her- bivory especially when plants were phosphate deficient. Total number of nematodes was higher with phosphate deficiency. Furthermore, non-significant peak values in soil respiration, total number of nematodes, and bacterial number were observed in phosphate deficient plants with high simulated herbivory. In phosphate amended plants, fast-growing protozoa and bacterial feeding nematodes decreased at high simulated herbivory. These results support the hypothesis that the plant regulates abundances of both AM and free-living rhizosphere organisms and thereby the amount of plant-available nutrients, according to demand via root exudation. Rhizobium spp. was significantly stimulated by phosphate addition but not affected by simulated herbivory. Metabolites produced by rhizosphere bacteria from plants exposed to high simulated herbivory in phosphate amended soil stimulated seed performance. Possible interactions between protozoa and nematodes in relation to production and composition of bacteria in the rhizosphere are discussed.     

Phosphorus acquisition from phytate depends on efficient bacterial grazing, irrespective of the mycorrhizal status of Pinus pinaster

Background and aims

Phosphorus from phytate, although constituting the main proportion of organic soil P, is unavailable to plants. Despite the well-known effects of rhizosphere trophic relationships on N mineralization, no work has been done yet on P mineralization. We hypothesized that the interactions between phytate-mineralizing bacteria, mycorrhizal fungi and bacterial grazer nematodes are able to improve plant P use from phytate.

Methods

We tested this hypothesis by growing Pinus pinaster seedlings in agar containing phytate as P source. The plants, whether or not ectomycorrhizal with the basidiomycete Hebeloma cylindrosporum, were grown alone or with a phytase-producing bacteria Bacillus subtilis and two bacterial-feeder nematodes, Rhabditis sp. and Acrobeloides sp. The bacteria and the nematodes were isolated from ectomycorrhizal roots and soil from P. pinaster plantations.

Results

Only the grazing of bacteria by nematodes enhanced plant P accumulation. Although plants increased the density of phytase-producing bacteria, these bacteria alone did not improve plant P nutrition. The seedlings, whether ectomycorrhizal or not, displayed a low capacity to use P from phytate.

Conclusions

In this experiment, the bacteria locked up the phosphorus, which was delivered to plant only by bacterial grazers like nematodes. Our results open an alternative route for better utilization of poorly available organic P by plants.     

Litter decomposition: a Russian matriochka doll

Litter is decomposed in a sequential process. In a concerted action animals and microorganisms break down complex organic matter to mineral products. Higher animals fragment and partially solubilize plant material. Subsequently, microorganisms (protozoa, fungi and bacteria) further degrade the organic matter to end products that cannot be metabolized further under the prevailing environmental conditions. During the process of decomposition some parts of the organic substrate and the excess energy are used to form new biomass. Some free organic intermediates may interact chemically to form relatively recalcitrant organic matter, such as humic substances. The degree of mineralization depends strongly on the type of organic matter in the litter and the physical and chemical conditions of the environment.     

The diversity and ecological significance of Protozoa

The unicellular eukaryotes are currently grouped in the kingdom Protista, together with their multicellular relatives. The inclusion of protozoa, algae and water moulds in a single taxon has resulted in nomenclatural problems, academic homelessness, and a reduction in their teaching. There are around 40 000 described protozoan protist species. Protozoa are principally grazers of bacteria, increasing mineralization and making nutrients more available to other organisms; most are aquatic, but they are also widespread animal parasites and symbionts. Their biomass, role in food chains, roles as mutualists and pathogens, and value as biomonitors are reviewed. To assess the role of protozoa in ecosystems more accurately, the current poor taxonomic standards in ecological work on protozoa must be improved. The manpower to respond to existing and new challenges in the field is declining as protozoology disappears from university courses and this problem needs to be addressed.     

不同土壤培肥措施下农田有机物分解的生态过程

通过在河北曲周实验站的田间试验,研究了4种不同土壤培肥措施条件下农田生态系统中几种主要土壤生物随有机物分解的变化规律、有机物的分解及其主要影响因素。研究结果表明：除土壤线虫外,其他几种主要的土壤生物的分布规律基本上是堆肥区〉原貌区〉对照区〉化肥区,与施入的有机物（小麦秸秆）的分解规律一致。在受人为扰动的堆肥区、化肥区和对照区土壤中,细菌占绝对优势,而在未开垦的原貌区中,真菌起着重要作用。 在有机物分解初期,土壤微生物能比较快地迁移到秸秆表面,秸秆表面的生物数量最多的是细菌,随着细菌的数量增加,原生动物数量亦呈现增加趋势,蚯蚓数量增多,而线虫的数量则减少。而有机物分解后期,真菌的数量逐渐减少,蚯蚓的数量也呈下降趋势,有机物的分解速度减慢。通过灰色关联度分析,9种外界因素（生物因素和环境因素）对小麦秸秆分解作用的相对重要程度排序：土壤温度（0．844）〉蚯蚓（0．777）〉真菌（0．764）〉全氮（0．754）〉线虫（0．753）〉有机质（0．742）〉细菌（O．738）〉原生动物（0．693）〉土壤含水量（0．661）,其中土壤温度和蚯蚓是影响土壤有机物分解的最重要的两个因素。     

Correlation of abundance of arbuscular mycorrhizal fungi, bacteria and saprophytic microfungi with soil carbon, nitrogen and phsophorus

A significant positive correlation between the concentration of CFU of soil saprophytic microfungi and total soil carbon content, organic matter (oxidizable carbon) and available phosphorus was observed in field collected soil samples. Concentration of CFU of culturable bacteria correlated negatively with soil organic matter. Specific length of hyphae of arbuscular mycorrhizal fungi in the soil correlated only with soil respiration rate. The results indicate that arbuscular mycorrhizal fungi are associated with soil microsites rich in some easily mineralizable fraction of soil, organic matter rather than with total or oxidizable organic carbon.     

The relative importance of the bacterial pathway and soil inorganic nitrogen increase across an extreme wood‐ash application gradient

Ash from combustion of biofuels, for example wood chips, is often deposited as waste, but due to its high content of essential plant nutrients and alkalinity, it has been proposed to recycle ash as a fertilizer and liming agent in biofuel production forest. However, current legislation sets rather strict limitations for wood‐ash application in biofuel production systems. The soil microfood web, that is microorganisms and their microfaunal grazers, protozoa and nematodes, is pivotal for essential ecosystem processes such as decomposition and plant nutrient release. Therefore, a thorough assessment of the impacts on microfood web structure and functioning must precede actions towards raising the currently allowed application rates. In a Danish Norway spruce plantation, we evaluate the impact of wood ash applied at dosages from 0 to the extreme case of 90 t ash ha^?1 on the microfood web, the bacterial community structure, soil content of inorganic nitrogen, organic matter, dissolved organic carbon and nitrogen. Using structural equation modelling (SEM), we disentangled the direct effect of the disturbance imposed by ash per se, the associated pH increase and changes in prey abundance on individual organism groups in the microfood web. The SEM showed that the pH rise was the main driver of increasing abundances of culturable heterotrophic bacteria with increasing ash doses, and via trophical transfer, this also manifested as higher abundances of bacterial grazers. Fungal‐feeding nematodes were unaffected by ash, whereas carnivorous/omnivorous nematodes decreased due to the direct effect of ash. Increasing ash doses enhanced the difference between bacterial communities of control plots and ash‐amended plots. The ash‐induced stimulation of culturable heterotrophic bacteria and bacterial grazers increased inorganic nitrogen availability at ash doses of 9 t ha^?1 and above. Hence, raised limits for ash application may potentially benefit tree growth via enhanced N mineralization activity of the soil food web.     

Plant metabolomics for plant chemical responses to belowground community change by climate change

General circulation models on global climate change predict increase in surface air temperature and changes in precipitation. Increases in air temperature (thus soil temperature) and altered precipitation are known to affect the species composition and function of soil microbial communities. Plant roots interact with diverse soil organisms such as bacteria, protozoa, fungi, nematodes, annelids and insects. Soil organisms show diverse interactions with plants (eg. competition, mutualism and parasitism) that may alter plant metabolism. Besides plant roots, various soil microbes such as bacteria and fungi can produce volatile organic compounds (VOCs), which can serve as infochemicals among soil organisms and plant roots. While the effects of climate change are likely to alter both soil communities and plant metabolism, it is equally probable that these changes will have cascading consequnces for grazers and subsequent food web components aboveground. Advances in plant metabolomics have made it possibile to track changes in plant metabolomes as they respond to biotic and abiotic environmental changes. Recent developments in analytical instrumentation and bioinformatics software have established metabolomics as an important research tool for studying ecological interactions between plants and other organisms. In this review, we will first summarize recent progress in plant metabolomics methodology and subsequently review recent studies of interactions between plants and soil organisms in relation to climate change issues.     

A comparison of nematode-capturing efficiencies of fiveDactylella species at four temperatures

Conclusions The data and observations presented indicate that the factors controlling the nematode-capturing efficiency of predaceous fungi are complex. There seems to be little correlation between the ability to capture nematodes and the optimum growth rate of the capturing fungus, but there may be some relation between the motility of a nematode and the ease with which it may be captured. Since soil contains many fungi and nematodes competing for nutrients in fairly restricted spheres such as soil particles, bits of organic matter, or the rhizospheres of growing plants, the influence of a nematode-supporting fungus such asPenicillium digitatum on a predaceous fungus may be greater than the direct effect of some nematode upon the predaceous fungus or vice versa.     

耕作方式对潮土土壤团聚体微生物群落结构的影响

为探究不同耕作方式对潮土土壤团聚体微生物群落结构和多样性的影响,采用磷脂脂肪酸(PLFA)法测定了土壤团聚体中微生物群落。试验设置4个耕作处理,分别为旋耕+秸秆还田(RT)、深耕+秸秆还田(DP)、深松+秸秆还田(SS)和免耕+秸秆还田(NT)。结果表明:与RT相比,DP处理显著提高了原状土壤和>5 mm粒级土壤团聚体中真菌PLFAs量和真菌/细菌,为真菌的繁殖提供了有利条件,有助于土壤有机质的贮存,提高了土壤生态系统的缓冲能力;提高了5～2 mm粒级土壤团聚体中细菌PLFAs量,降低了土壤革兰氏阳性菌/革兰氏阴性菌,改善了土壤营养状况;提高了<0.25 mm粒级土壤团聚体中微生物丰富度指数。总的来说,深耕+秸秆还田(DP)对土壤团聚体细菌和真菌生物量有一定的提高作用,并且在一定程度上改善了土壤团聚体微生物群落结构,有利于增加土壤固碳能力和保持土壤微生物多样性。冗余分析结果表明,土壤团聚体总PLFAs量、细菌、革兰氏阴性菌和放线菌PLFAs量与土壤有机碳相关性较强,革兰氏阳性菌PLFAs量与总氮相关性较强。各处理较大粒级土壤团聚体微生物群落主要受碳氮比、含水量、pH值和团聚体质量分数的影响,较小粒级土壤团聚体微生物群落则主要受土壤有机碳和总氮的影响。     

Nematode Indicators of Organic Enrichment

The organisms of the soil food web, dependent on resources from plants or on amendment from other sources, respond characteristically to enrichment of their environment by organic matter. Primary consumers of the incoming substrate, including bacteria, fungi, plant-feeding nematodes, annelids, and some microarthropods, are entry-level indicators of enrichment. However, the quantification of abundance and biomass of this diverse group, as an indicator of resource status, requires a plethora of extraction and assessment techniques. Soluble organic compounds are absorbed by bacteria and fungi, while fungi also degrade more recalcitrant sources. These organisms are potential indicators of the nature of incoming substrate, but current methods of biomass determination do not reliably indicate their community composition. Guilds of nematodes that feed on bacteria (e.g., Rhabditidae, Panagrolaimidae) and fungi (e.g., Aphelenchidae, Aphelenchoididae) are responsive to changes in abundance of their food. Through direct herbivory, plant-feeding nematodes (e.g., many species of Tylenchina) also contribute to food web resources. Thus, analysis of the nematode community of a single sample provides indication of carbon flow through an important herbivore channel and through channels mediated by bacteria and fungi. Some nematode guilds are more responsive than others to resource enrichment. Generally, those bacterivores with short lifecycles and high reproductive potential (e.g., Rhabditidae) most closely mirror the bloom of bacteria or respond most rapidly to active plant growth. The feeding habits of some groups remain unclear. For example, nematodes of the Tylenchidae may constitute 30% or more of the individuals in a soil sample; further study is necessary to determine which resource channels they portray and the appropriate level of taxonomic resolution for this group. A graphic representation of the relative biomass of bacterivorous, fungivorous, and herbivorous nematodes provides a useful tool for assessing the importance of the bacterial, fungal, and plant resource channels in an extant food web.     

Organic matter–microorganism–plant in soil bioremediation: a synergic approach

Bioremediation is a natural process, which relies on bacteria, fungi, and plants to degrade, break down, transform, and/or essentially remove contaminants, ensuring the conservation of the ecosystem biophysical properties. Since microorganisms are the former agents for the degradation of organic contaminants in soil, the application of organic matter (such as compost, sewage sludge, etc.), which increases microbial density and also provides nutrients and readily degradable organic matter (bioenhancement–bioaugmentation) can be considered useful to accelerate the contaminant degradation. Moreover, the organic matter addition, by means of the increase of cation exchange capacity, soil porosity and water-holding capacity, enhances the soil health and provides a medium satisfactory for microorganism activity. Plants have been also recently used in soil reclamation strategy both for their ability to uptake, transform, and store the contaminants, and to promote the degradation of organic contaminants by microbes at rhizosphere level. It is widely recognized that plant, through organic materials, nutrients and oxygen supply, produces a rich microenvironment capable of promoting microbial proliferation and activity.     

Effect of Introduced Pseudomonas fluorescens Strains on Soil Nematode and Protozoan Populations in the Rhizosphere of Wheat and Pea

Abstract Previous studies have shown that inoculation of pea seeds with Pseudomonas fluorescens strains F113lacZY or F113G22 increased mineralization of organic nitrogen in the rhizosphere. In contrast, inoculation of the same strains onto wheat seeds reduced mineralization of N from organic residues incorporated into soil. In the present study, we report on a likely explanation of this phenomenon, which appears to be governed by the effect of plant-microbe interactions on bacterial-feeding nematodes and protozoa. In soil microcosm tests, inoculation of pea seeds with Pseudomonas fluorescens strains F113lacZY or F113G22 resulted in an increase in the number of nematodes and protozoa in the rhizosphere as compared to noninoculated controls. This trend was repeated using a model sand system into which the bacteriophagous nematode Caenorhabditis elegans was introduced. It was subsequently found that non-inoculated germinating pea seeds exerted a nematicidal effect on C. elegans, which was remedied by inoculation with either strain F113lacZY or F113G22. This suggests that nematicidal compounds released by the germinating pea seeds were metabolized by the microbial inoculants before they affected nematode populations in the spermosphere or rhizosphere of pea. In contrast, inoculation of wheat plants resulted in significantly lower nematode populations in the rhizosphere, whereas protozoan numbers were unaffected. No nematicidal effects of inoculated or noninoculated wheat seeds could be found, suggesting that microfaunal populations were affected at a later stage during plant growth. Because of their key roles in accelerating the turnover of microbially immobilized N and organic matter, plants that support a larger microfaunal population are likely to benefit from a higher availability of inorganic nitrogen. Therefore, an understanding of plant-microbe interactions and their effects on soil microfaunal populations is essential in order to assess the effects of microbial inocula on plant mineral nutrition. Received: 27 May 1999; Accepted: 15 July 1999; Online Publication: 17 December 1999     

Six years of in situ CO2 enrichment evoke changes in soil structure and soil biota of nutrient-poor grassland

Nutrient‐poor grassland on a silty clay loam overlying calcareous debris was exposed to elevated CO₂ for six growing seasons. The CO₂ exchange and productivity were persistently increased throughout the experiment, suggesting increases in soil C inputs. At the same time, elevated CO₂ lead to increased soil moisture due to reduced evapotransporation. Measurements related to soil microflora did not indicate increased soil C fluxes under elevated CO₂. Microbial biomass, soil basal respiration, and the metabolic quotient for CO₂ (qCO₂) were not altered significantly. PLFA analysis indicated no significant shift in the ratio of fungi to bacteria. 0.5 m KCl extractable organic C and N, indicators of changed DOC and DON concentrations, also remained unaltered. Microbial grazer populations (protozoa, bacterivorous and fungivorous nematodes, acari and collembola) and root feeding nematodes were not affected by elevated CO₂. However, total nematode numbers averaged slightly lower under elevated CO₂ (?16%, ns) and nematode mass was significantly reduced (?43%, P = 0.06). This reduction reflected a reduction in large‐diameter nematodes classified as omnivorous and predacious. Elevated CO₂ resulted in a shift towards smaller aggregate sizes at both micro‐ and macro‐aggregate scales; this was caused by higher soil moisture under elevated CO₂. Reduced aggregate sizes result in reduced pore neck diameters. Locomotion of large‐diameter nematodes depends on the presence of large enough pores; the reduction in aggregate sizes under elevated CO₂ may therefore account for the decrease in large nematodes. These animals are relatively high up the soil food web; this decline could therefore trigger top‐down effects on the soil food web. The CO₂ enrichment also affected the nitrogen cycle. The N stocks in living plants and surface litter increased at elevated CO₂, but N in soil organic matter and microbes remained unaltered. Nitrogen mineralization increased markedly, but microbial N did not differ between CO₂ treatments, indicating that net N immobilization rates were unaltered. In summary, this study did not provide evidence that soils and soil microbial communities are affected by increased soil C inputs under elevated CO₂. On the contrary, available data (¹³C tracer data, minirhizotron observations, root ingrowth cores) suggests that soil C inputs did not increase substantially. However, we provide first evidence that elevated CO₂ can reduce soil aggregation at the scale from µ m to mm scale, and that this can affect soil microfaunal populations.     

Aphid effects on rhizosphere microorganisms and microfauna depend more on barley growth phase than on soil fertilization

This paper gives the first reports on aphid effects on rhizosphere organisms as influenced by soil nutrient status and plant development. Barley plants grown in pots fertilized with N but without P (N), with N and P (NP), or not fertilized (0) were sampled in the early growth phase (day 25), 1 week before and 1 week after spike emergence. Aphids were added 16 days before sampling was carried out. In a separate experiment belowground respiration was measured on N and NP fertilized plant–soil systems with aphid treatments comparable to the first experiment. Aphids reduced numbers of rhizosphere bacteria and fungal feeding nematodes 1 week before spike emergence. Before spike emergence, aphids reduced belowground respiration in NP treatments. These findings strongly indicate that aphids reduced allocation of photoassimilates to roots and deposition of root exudates in the growth phase of the plant. Contrary to this, 1 week after spike emergence numbers of bacteria, fungal feeding nematodes and Protozoa were higher in rhizospheres of plants subjected to aphids probably because aphids enhanced root mortality and root decomposition. Protozoa and bacterial feeding nematodes were stimulated at different experimental conditions with nematodes being the dominant bacterial grazers at N fertilization and Protozoa in the NP treatment before spike emergence.     

Coordination of aboveground and belowground responses to local‐scale soil fertility differences between two contrasting Jamaican rain forest types

There is growing interest in understanding how declining soil fertility in the prolonged absence of major disturbance drives ecological processes, or ‘ecosystem retrogression’. However, there are few well characterized study systems for exploring this phenomenon in the tropics, despite tropics occupying over 40% of the Earth's terrestrial surface. We studied two types of montane rain forest in the Blue Mountains of Jamaica that represent distinct stages in ecosystem development, i.e. an earlier stage with shallow organic matter and a late stage with deep organic matter (hereafter ‘mull’ and ‘mor’ stages). We characterized responses of soil fertility and plant, soil microbial and nematode communities to the transition from mull to mor and whether these responses were coupled. For soil abiotic properties, we found this transition led to lower amounts of both nitrogen (N) and phosphorus (P) and an enhanced N to P ratio. This led to shorter‐statured and less diverse forest, and convergence of tree species composition among plots. At the whole community (but not individual species) level foliar and litter N and P diminished from mull to mor, while foliar N to P and resorption efficiency of P relative to N increased, indicating increasing P relative to N limitation. We also found impairment of soil microbes (but not nematodes) and an increasing role of fungi relative to bacteria during the transition. Our results show that retrogression phenomena involving increasing nutrient (notably P) limitation can be important drivers in tropical systems, and are likely to involve aboveground–belowground feedbacks whereby plants produce litter of diminishing quality, impairing soil microbial processes and thus reducing the supply of nutrients from the soil for plant growth. Such feedbacks between plants and the soil, mediated by plant litter and organic matter quality, may serve as major though often overlooked drivers of long term environmental change.