Multiple factors influence the role of arbuscular mycorrhizal fungi in soil aggregation—a meta-analysis

Background and aims

Soil aggregation is a crucial aspect of ecosystem functioning in terrestrial ecosystems. Arbuscular mycorrhizal fungi (AMF) play a key role in soil aggregate formation and stabilization. Here we quantitatively analyzed the importance of experimental settings as well as biotic and abiotic factors for the effectiveness of AMF to stabilize soil macroaggregates.

Methods

We gathered 35 studies on AMF and soil aggregation and tested 13 predictor variables for their relevance with a boosted regression tree analysis and performed a meta-analysis, fitting individual random effects models for each variable.

Results and conclusions

The overall mean effect of inoculation with AMF on soil aggregation was positive and predictor variable means were all in the range of beneficial effects. Pot studies and studies with sterilized sandy soil, near neutral soil pH, a pot size smaller than 2.5 kg and a duration between 2.2 and 5 months were more likely to result in stronger effects of AMF on soil aggregation than experiments in the field, with non-sterilized or fine textured soil or an acidic pH. This is the first study to quantitatively show that the effect of AMF inoculation on soil aggregation is positive and context dependent. Our findings can help to improve the use of this important ecosystem process, e.g. for inoculum application in restoration sites.     

Mechanisms linking plant community properties to soil aggregate stability in an experimental grassland plant diversity gradient

Background and aims

Soil aggregate stability depends on plant community properties, such as functional group composition, diversity and biomass production. However, little is known about the relative importance of these drivers and the role of soil organisms in mediating plant community effects.

Methods

We studied soil aggregate stability in an experimental grassland plant diversity gradient and considered several explanatory variables to mechanistically explain effects of plant diversity and plant functional group composition. Three soil aggregate stability measures (slaking, mechanical breakdown and microcracking) were considered in path analyses.

Results

Soil aggregate stability increased significantly from monocultures to plant species mixtures and in the presence of grasses, while it decreased in the presence of legumes, though effects differed somewhat between soil aggregate stability measures. Using path analysis plant community effects could be explained by variations in root biomass, soil microbial biomass, soil organic carbon concentrations (all positive relationships), and earthworm biomass (negative relationship with mechanical breakdown).

Conclusions

The present study identified important drivers of plant community effects on soil aggregate stability. The effects of root biomass, soil microbial biomass, and soil organic carbon concentrations were largely consistent across plant diversity levels suggesting that the mechanisms identified are of general relevance.     

Drying and wetting in saline and saline-sodic soils—effects on microbial activity, biomass and dissolved organic carbon

Aims

There are few studies on the interactive effect of salinity and sodicity in soils exposed to drying and wetting cycles. We conducted a study to assess the impact of multiple drying and wetting on microbial respiration, dissolved organic carbon and microbial biomass in saline and saline-sodic soils.

Methods

Different levels of salinity (EC_1:5 1.0 or 2.5) and sodicity (SAR?<?3 or 20) were induced by adding NaCl and CaCl₂ to a non-saline/non-sodic soil. Finely ground wheat straw residue was added at 20?g?kg^?1 as substrate to stimulate microbial activity. The constant moist (CM) treatment was kept at optimum moisture content for the length of the experiment. The drying and rewetting (DW) treatments consisted of 1 to 3 DW cycles; each DW cycle consisted of 1?week drying after which they were rewet to optimum moisture and then maintained moist for 1?week.

Results

Drying reduced respiration more strongly at EC2.5 than with EC1.0. Rewetting of dry soils produced a flush in respiration which was greatest in the soils without salt addition and smallest at high salinity (EC2.5) suggesting better substrate utilisation by microbes in soils without added salts. After three DW events, cumulative respiration was significantly increased by DW compared to CM, being 24% higher at EC1.0 and 16% higher at EC2.5 indicating that high respiration rates after rewetting may compensate for the low respiration rates during the dry phase. The respiration rate per unit MBC was lower at EC2.5 than at EC1.0. Further, the size of the flush in respiration upon rewetting decreased with each ensuing DW cycle being 50–70% lower in the third DW cycle than the first.

Conclusions

Both salinity and sodicity alter the effect of drying and rewetting on soil carbon dynamics compared to non-saline soils.     

Signalling mechanisms involved in the response of two varieties of Humulus lupulus L. to soil drying: I. changes in xylem sap pH and the concentrations of abscisic acid and anions

Background and aims

Soil drying leads to the generation of chemical signals in plants that regulate water use via control of the stomatal aperture. The aim of our work was to identify the presence and identity of potential chemical signals, their dynamics, and their relationship with transpiration rate during soil drying in hop (Humulus lupulus (L.)) plants.

Methods

We used pressure chamber technique for measurement of shoot water potential and collection of shoot xylem sap. We analyzed concentrations of abscisic acid (ABA), nitrate, phosphate, sulphate and malate in sap and also the rate of whole plant transpiration.

Results

Transpiration rate decreased prior to changes in shoot water potential. The concentration of ABA in xylem sap continuously increased from early to later stages of water stress, whereas in leaves it increased only at later stages. Shoot sap pH increased simultaneously with the decrease of transpiration rate. Xylem sap alkalization was in some cases accompanied by a decrease in nitrate concentration and an increase in malate concentration. Concentration of sulphate increased in xylem sap during drying and sulphate in combination with a higher ABA concentration enhanced stomatal closure.

Conclusions

Several early chemical signals appear in sap of hop plants during soil drying and their impact on transpiration may vary according to the stage of soil drying.     

Responses of CO2 efflux from an alpine meadow soil on the Qinghai Tibetan Plateau to multi-form and low-level N addition

Aims

To assess the effects of atmospheric N deposition on the C budget of an alpine meadow ecosystem on the Qinghai–Tibetan Plateau, it is necessary to explore the responses of soil-atmosphere carbon dioxide (CO₂) exchange to N addition.

Methods

Based on a multi-form, low-level N addition experiment, soil CO₂ effluxes were monitored weekly using the static chamber and gas chromatograph technique. Soil variables and aboveground biomass were measured monthly to examine the key driving factors of soil CO₂ efflux.

Results

The results showed that low-level N input tended to decrease soil moisture, whereas medium-level N input maintained soil moisture. Three-year N additions slightly increased soil inorganic N pools, especially the soil NH ₄ ⁺ -N pool. N applications significantly increased aboveground biomass and soil CO₂ efflux; moreover, this effect was more significant from NH ₄ ⁺ -N than from NO ₃ ^? -N fertilizer. In addition, the soil CO₂ efflux was mainly driven by soil temperature, followed by aboveground biomass and NH ₄ ⁺ -N pool.

Conclusions

These results suggest that chronic atmospheric N deposition will stimulate soil CO₂ efflux in the alpine meadow on the Qinghai–Tibetan Plateau by increasing available N content and promoting plant growth.     

Degradation of moist soil aggregates by rapid temperature rise under low intensity fire

Background and aims

Soil structure degradation by fire is usually attributed to qualitative and quantitative change of organic and inorganic binding agents, especially in high severity burns (>300 °C) that last for prolonged periods (> 1 hour). In contrast, controlled burns are typically managed to be low in intensity and severity. Such burns are considered benign to soil structural stability because organic matter and inorganic binding agents (e.g., gypsum) are relatively stable at such low temperatures. Recent observations at a controlled burn site in the eastern Great Basin (Nevada) showed soil aggregate breakdown found in shrub canopies where soil temperatures briefly exceeded 300 °C as well as interspaces between shrubs, where the temperatures were likely lower than beneath shrubs because of less surface biomass. These alterations cannot be explained in terms of thermal alteration of binding agents. This study was designed to test whether pressure created by rapidly vaporized pore water can cause aggregate breakdown.

Methods

We subjected three different sizes of aggregates (0.25–1, 1–2 and 2–4 mm) of soils derived from the eastern Great Basin burn site as well as from a forest and urban garden in California to rapid and slow (3 °C/min) heating rates. These treatments were conducted at 5 peak temperatures (75, 100, 125, 150 and 175 °C).

Results

Post-burn water stability of the aggregates showed that rapid heating rate caused more pronounced degradation of aggregate stability than slow heating. Moreover, the heating-rate dependent structural degradation increased with peak temperature. For the majority of the aggregates, the effect also increased with initial water content. In all the soils tested, there was no preferential loss of organic matter in the rapid-heating treatment that can explain the observed enhanced breakdown of aggregates.

Conclusions

Our observations indicate that soil structural degradation under low-intensity fire occurs as a result of mechanical stresses extorted by rapidly escaping steam from soil pores under rapid heating rate.     

Root plasticity maintains growth of temperate grassland species under pulsed water supply

Background and aims

The frequency of rain is predicted to change in high latitude areas with more precipitation in heavy, intense events interspersed by longer dry periods. These changes will modify soil drying cycles with unknown consequences for plant performance of temperate species.

Methods

We studied plant growth and root traits of juveniles of four grasses and four dicots growing in a greenhouse, when supplying the same total amount of water given either regular every other day or pulsed once a week.

Results

Pulsed water supply replenished soil moisture immediately after watering, but caused substantial drought stress at the end of the watering cycle, whereas regular watering caused more moderate but consistent drought. Grasses had lower water use efficiency in the pulsed watering compared to regular watering, whereas dicots showed no difference. Both grasses and dicots developed thinner roots, thus higher specific root length, and greater root length in the pulsed watering. Growth of dicots was slightly increased under pulsed watering.

Conclusions

Temperate species coped with pulsed water supply by eliciting two responses: i) persistent shoot growth, most likely by maximizing growth at peaks of soil moisture, thus compensating for slower growth during drought periods; ii) plasticity of root traits related to increased resource uptake. Both responses likely account for subtle improvement of growth under changed water supply conditions.     

Quantifying the effect of soil moisture content on segmenting root system architecture in X-ray computed tomography images

Aims

A commonly accepted challenge when visualising plant roots in X-ray micro Computed Tomography (μCT) images is the similar X-ray attenuation of plant roots and soil phases. Soil moisture content remains a recognised, yet currently uncharacterised source of segmentation error. This work sought to quantify the effect of soil moisture content on the ability to segment roots from soil in μCT images.

Methods

Rice (Oryza sativa) plants grown in contrasting soils (loamy sand and clay loam) were μCT scanned daily for nine days whilst drying from saturation. Root volumes were segmented from μCT images and compared with volumes derived by root washing.

Results

At saturation the overlapping attenuation values of root material, water-filled soil pores and soil organic matter significantly hindered segmentation. However, in dry soil (ca. six days of drying post-saturation) the air-filled pores increased image noise adjacent to roots and impeded accurate visualisation of root material. The root volume was most accurately segmented at field capacity.

Conclusions

Root volumes can be accurately segmented from μCT images of undisturbed soil without compromising the growth requirements of the plant providing soil moisture content is kept at field capacity. We propose all future studies in this area should consider the error associated with scanning at different soil moisture contents.     

Does biochar influence soil physical properties and soil water availability?

Aims

This study aims to (i) determine the effects of incorporating 47 Mg ha^?1 acacia green waste biochar on soil physical properties and water relations, and (ii) to explore the different mechanisms by which biochar influences soil porosity.

Methods

The pore size distribution of the biochar was determined by scanning electron microscope and mercury porosimetry. Soil physical properties and water relations were determined by in situ tension infiltrometers, desorption and evaporative flux on intact cores, pressure chamber analysis at ?1,500 kPa, and wet aggregate sieving.

Results

Thirty months after incorporation, biochar application had no significant effect on soil moisture content, drainable porosity between –1.0 and ?10 kPa, field capacity, plant available water capacity, the van Genuchten soil water retention parameters, aggregate stability, nor the permanent wilting point. However, the biochar-amended soil had significantly higher near-saturated hydraulic conductivity, soil water content at ?0.1 kPa, and significantly lower bulk density than the unamended control. Differences were attributed to the formation of large macropores (>1,200 μm) resulting from greater earthworm burrowing in the biochar-amended soil.

Conclusion

We found no evidence to suggest application of biochar influenced soil porosity by either direct pore contribution, creation of accommodation pores, or improved aggregate stability.     

Changes in microbial activities and biomasses over a forest floor gradient in C-to-N ratio

Background and aims

Under chronically elevated N deposition, N retention mainly occur at high soil C-to-N ratio. This may be mediated through soil microbes, such as ectomycorrhizal (EM) fungi, saprotrophic fungi and bacteria, and the aim of this study was to evaluate the relationship between soil microbes and forest floor C-to-N ratios.

Methods

Soil samples from 33 Norway spruce (Picea abies (L.) H. Karst) forests in Denmark and southern Sweden in a forest floor C-to-N ratio gradient (ranging from 14 to 35) were analysed regarding the content of phospholipid fatty acids (PLFAs) to estimate their soil microbial community composition and the relative biomasses of different microbial groups. The relation of EM biomass to total fungal biomass was estimated as the loss of the fungal PLFA 18:2ω6,9 during incubation of soils and the production of EM mycelia was estimated using fungal in-growth mesh bags. The soil microbial variables were correlated to forest floor C-to-N ratio, NO ₃ ^- leaching, soil pH and stand age.

Results

Fungal proportions of microbial biomass, EM to total fungi and EM mycelial production were all positively related to C-to-N ratio, while NO ₃ ^- leaching was negatively related to C-to-N ratio.

Conclusions

Both EM and saprotrophic fungi change with forest floor C-to-N ratios and appear to play a central role in N retention in forest soil. A better understanding of the mechanisms behind this process may be revealed if the role of recalcitrant fungal metabolites for N retention (and soil C sequestration) can be identified. Research along this line deserves further studies.     

Mechanical properties and soil stability affected by fertilizer treatments for an Ultisol in subtropical China

Background and aims

Soil mechanical properties are crucial for plant growth, soil erosion, tillage and traffic. The soil mechanical properties and stability of an Ultisol were determined in a 13-year fertilization experiment in subtropical China. The effect of organic matter on soil structure was also evaluated.

Methods

The treatments include: unfertilized, mineral fertilized, mineral mixed with straw, and animal manure. Bulk soil strengths (shear strength and penetration resistance) were tested in field. Aggregate strengths (penetration resistance and tensile strength), water stability, organic carbon (OC), hot-water-extractable carbohydrate (HWEC) and some related factors were determined in laboratory.

Results

Fertilizer increases aggregate penetration resistance, tensile strength, water stability and organic matter content in cultivated horizon (0–15 cm depth), especially at the 0–5 cm layer. OC and HWEC showed significantly regression relationships with aggregate water stability, porosity and water repellency for the whole soil profile (0–40 cm). Aggregate strengths also increased with OC and HWEC in cultivated horizon.

Conclusions

Animal manure is most effective in improving soil structure while aggregate water stability is the most sensitive index of fertilizer management. Soil organic matter is a strong determinant of soil structure. Both OC and HWEC are good indicators of surface aggregate strengths and stability, and thus, soil quality in this region.     

Comparison of common bean (Phaseolus vulgaris L.) genotypes for nitrogen fixation tolerance to soil drying

Aims

Common bean is a major source of protein for many people worldwide. However, the crop is often subjected to drought conditions and its advantage in undertaking symbiotic nitrogen fixation can be severely decreased. The primary objective of this study was to compare the resistance of nitrogen fixation of 12 selected genotypes to soil drying.

Methods

Twelve common bean genotypes of diverse genetic background were compared. Plants were grown in pots and subjected to soil drying over about 2 weeks. Nitrogen fixation was measured daily using a flow-through acetylene reduction technique. The plants were exposed to acetylene for only a short time period allowing repeated measures. The acetylene reduction rate of plants on drying soil was normalized against the rates measured for well-watered plants.

Results

Substantial variability was identified among genotypes in the threshold soil water content at which nitrogen fixation was observed to decrease. Genotypes SER 16, SXB 412, NCB 226, and Calima were found to have the greatest delay in their decrease in nitrogen fixation rates based on soil water content. These four genotypes expressed substantial tolerance of nitrogen fixation to soil drying. These experiments also resulted in data on the threshold soil water contents at which transpiration rates decreased. A decrease in transpiration rates at high soil water contents is potentially advantageous since it allows soil water conservation for use as the severity of the drought increases. There was a general trend of those genotypes with sustained nitrogen fixation rates to low soil water contents also expressing decreased transpiration rates at high soil water contents.

Conclusions

This study identified genetic variation among common bean genotypes in their response of nitrogen fixation and transpiration to soil drying. Five genotypes (SER 16, SXB 412, NCB 226, Calima, and SEA 5) expressed the desired traits for water-limited conditions, which might be exploited in breeding efforts.     

Coupled soil-availability and tree-limitation nutritional shifts induced by N deposition: insights from N to P relationships in Abies pinsapo forests

Background and aims

Interacting effects of atmospheric N deposition on the degree to which tree demand for other nutrients is met by soil supply has seldom been explored in Mediterranean-type ecosystems. We hypothesized that patterns for the relative availability of N and P in soils will be matched by variations in process rates related to soil organic P cycling and by shifts from N to P limitation of tree growth.

Methods

We examined N/P relationships in Mediterranean-fir (Abies pinsapo) forests from two nearby regions differing in N deposition levels.

Results

N pools and transformation rates and the contribution of organic fractions to the labile P pool in soils showed increasing trends toward the pollution source. Phosphomonoesterase activity (PME) in bulk soils, root PME per unit biomass (but not per unit soil volume) and biomass accumulation in P-fertilized root-in-growth cores incubated in situ were also the highest at the sites receiving elevated N deposition, indicating P limitation. In contrast, forest stands in the region farther from the pollutant source were N-limited (preferential root growth in N-rich soil microsites) and showed lower PME activities and higher total fine root biomass.

Conclusions

In the forests under elevated N deposition, higher values for an overall indicator of soil N status matched with indications of an accelerated soil organic P subcycle and P-limitation of tree growth.     

The influence of 3-years of warming and N-deposition on ecosystem dynamics is small compared to past land use in subalpine meadows

Background and aims

Drivers of ecosystem dynamics that are under human influence range from local, land-management decisions to global processes such as warming temperatures and N deposition. The goal of this study was to understand how multiple, potentially interacting factors influence net primary production, N mineralization, and water and soil CO₂ fluxes.

Methods

Here I report on a three-year experiment that manipulated air temperature using ITEX passive warming cones and N deposition in a mountain meadow ecosystems that were historically grazed or protected from grazing.

Results

The strongest and most consistent effect was due to the legacy of grazing, with previously grazed sites having lower primary production, lower soil respiration rates, lower soil moisture, and lower soil C and N stocks than historically ungrazed sites. Warming increased soil respiration, but the effect was transient, and decreased over the 3-year study. Nitrogen addition increased primary production in the second and third year of the experiment but had no significant effect on soil respiration. The effect of historical grazing on primary production was approximately double the effect of N addition. Temperature and N deposition rarely interacted except for increasing N availability during the warm, wet growing season of 2004.

Conclusions

These findings indicate that the legacies of land use, with their influence on plant community composition and hydrologic processes, are locally more important than short-term step changes in temperature and nutrient availability.     

Modelling the impact of heterogeneous rootzone water distribution on the regulation of transpiration by hormone transport and/or hydraulic pressures

Aims

A simulation model to demonstrate that soil water potential can regulate transpiration, by influencing leaf water potential and/or inducing root production of chemical signals that are transported to the leaves.

Methods

Signalling impacts on the relationship between soil water potential and transpiration were simulated by coupling a 3D model for water flow in soil, into and through roots (Javaux et al. 2008) with a model for xylem transport of chemicals (produced as a function of local root water potential). Stomatal conductance was regulated by simulated leaf water potential (H) and/or foliar chemical signal concentrations (C; H?+?C). Split-root experiments were simulated by varying transpiration demands and irrigation placement.

Results

While regulation of stomatal conductance by chemical transport was unstable and oscillatory, simulated transpiration over time and root water uptake from the two soil compartments were similar for both H and H?+?C regulation. Increased stomatal sensitivity more strongly decreased transpiration, and decreased threshold root water potential (below which a chemical signal is produced) delayed transpiration reduction.

Conclusions

Although simulations with H?+?C regulation qualitatively reproduced transpiration of plants exposed to partial rootzone drying (PRD), long-term effects seemed negligible. Moreover, most transpiration responses to PRD could be explained by hydraulic signalling alone.     

Carbon allocation in Larrea tridentata plant-soil systems as affected by elevated soil moisture and N availability

Background and Aims

Global change will likely express itself in southwestern United States arid lands through changes in amounts and timing of precipitation in response to elevated CO₂ concentrations. In addition, increased nitrogen (N) deposition may occur due to increased urban development. This study addressed the effects of water and N availability on C allocation in arid land soil-plant systems.

Methods

Columns filled with Mojave Desert topsoil containing Larrea tridentata seedlings with two treatment levels each of N and soil moisture were labeled by exposure to ¹³C-enriched CO₂.

Results

Increased soil moisture increased plant biomass, total ¹³C uptake, ¹³C levels in leaves, soil organic matter, and soil respiration, decreased relative C allocation to stems but increased allocation to soil organic matter. Increased soil N availability increased N uptake but decreased C allocation to soil respiration presumably due to decreased substrate supply for microbes. There was no detectable label in carbonate C, suggesting that this pool does not significantly contribute to ecosystem C fluxes.

Conclusions

Our study indicates that increased water availability causes increased C uptake with increased C allocation to soil organic matter in Larrea tridentata-dominated communities while increased N deposition will have a minimal impact on C sequestration.     

Effects of experimental warming and nitrogen fertilization on soil microbial communities and processes of two subalpine coniferous species in Eastern Tibetan Plateau,China

Aim

This study aimed at predicting how sub-alpine coniferous ecosystems respond to global changes in the Eastern Tibetan Plateau by understanding soil microbial communities and activities, as well as variation in the quality and quantity of soil organic matter.

Methods

An experiment was conducted to examine soil microbial communities and their related soil processes in rhizospheric soil of two coniferous species that were exposed to two levels of temperature (unwarmed and infrared heater warming) and two levels of nitrogen (unfertilized and 25 g N m^?2 a^?1) from April 2007.

Results

Four-year night warming alone slightly affected the phospholipid fatty acid contents of the microbial community. However, the combination of nitrogen addition and soil warming significantly affected soil microbial composition while reducing the biomass of major microbial groups and the activities of most enzymes, especially in Abies faxoniana plots. The combination of warming and nitrogen addition increased soil labile C and N pools in Picea asperata plots and was beneficial for soil recalcitrant C, as well as for labile and total C and N pools in A. faxoniana plots.

Conclusion

Results indicated that future warming will slightly affect soil microbial communities and their related soil processes. However, warming combined with high nitrogen deposition will significantly constrain soil microbial biomass and enzyme activities, consequently increasing soil C and N pools in sub-alpine coniferous forests of this region.     

Restoration of wetlands: successes and failures on scalds comprising an iron oxide clogged layer with areas of acid sulfate soils

Background and aims

We evaluated the influence of plant species and life forms on soil aggregate distribution among size-classes, total macroaggregate mass and aggregate mean weight diameter (MWD), and examined how specific root traits were related to these aggregation variables.

Methods

We analyzed the soil attached to the roots (i.e., rhizospheric soil) under 13 Mediterranean species grown in monocultures in a common garden experiment for four years, and compared it to a bare soil. The mass distribution of aggregates in six size-classes and aggregate MWD were calculated, both on a rhizospheric soil and root biomass basis.

Results

Compared to bare soil, macroaggregate mass increased by an average of 13% in the presence of plants, with a strong effect of species and life forms (both P < 0.0001); some species such as Sanguisorba minor showing increases of up to ~40%. Although the soil under graminoids had a greater macroaggregate mass, their MWD was lower than under non-woody dicots. Large (2000–1000 μm) and intermediate (1000–500 μm) macroaggregate mass increased with root mass and length density and decreased with root lignin concentration, while very large macroaggregate (6000–2000 μm) mass and the MWD increased with root soluble compound concentration.

Conclusions

Species and life forms differently influenced the distribution of macroaggregates among size-classes and aggregate MWD. Easily-decomposable roots with traits related to resource acquisition (i.e., high fine root length, high water-soluble compound concentration) are more favorable for the development of water-stable macroaggregates than roots traits related to resource conservation (high lignin concentration, thick roots).

     

Regulation of aggregate size and pattern by adenosine and caffeine in cellular slime molds

Background

Multicellularity in cellular slime molds is achieved by aggregation of several hundreds to thousands of cells. In the model slime mold Dictyostelium discoideum, adenosine is known to increase the aggregate size and its antagonist caffeine reduces the aggregate size. However, it is not clear if the actions of adenosine and caffeine are evolutionarily conserved among other slime molds known to use structurally unrelated chemoattractants. We have examined how the known factors affecting aggregate size are modulated by adenosine and caffeine.

Result

Adenosine and caffeine induced the formation of large and small aggregates respectively, in evolutionarily distinct slime molds known to use diverse chemoattractants for their aggregation. Due to its genetic tractability, we chose D. discoideum to further investigate the factors affecting aggregate size. The changes in aggregate size are caused by the effect of the compounds on several parameters such as cell number and size, cell-cell adhesion, cAMP signal relay and cell counting mechanisms. While some of the effects of these two compounds are opposite to each other, interestingly, both compounds increase the intracellular glucose level and strengthen cell-cell adhesion. These compounds also inhibit the synthesis of cAMP phosphodiesterase (PdsA), weakening the relay of extracellular cAMP signal. Adenosine as well as caffeine rescue mutants impaired in stream formation (pde4 ^- and pdiA ^- ) and colony size (smlA ^- and ctnA ^- ) and restore their parental aggregate size.

Conclusion

Adenosine increased the cell division timings thereby making large number of cells available for aggregation and also it marginally increased the cell size contributing to large aggregate size. Reduced cell division rates and decreased cell size in the presence of caffeine makes the aggregates smaller than controls. Both the compounds altered the speed of the chemotactic amoebae causing a variation in aggregate size. Our data strongly suggests that cytosolic glucose and extracellular cAMP levels are the other major determinants regulating aggregate size and pattern. Importantly, the aggregation process is conserved among different lineages of cellular slime molds despite using unrelated signalling molecules for aggregation.