The impact of weed diversity on insect population dynamics and crop yield in collards, Brassica oleraceae (Brassicaceae)

Vegetational diversity within agricultural fields is often suggested as a means to reduce insect herbivore populations and to increase their natural enemies. In this paper we compare population densities of herbivores, predators, and parasitoids on collards in monocultures and on collards interplanted with two different groups of weeds, one with weed species from the same plant family as the collards (Brassicaceae) and one with weed species from unrelated plant families (non-Brassicaceae). The collards in the Brassicaceae weed polyculture had higher densities (number of herbivores/mean leaf area (cm²) per plant) of specialist herbivores than collards in the non-Brassicaceae weed polyculture and in collard monoculture. The “resource concentration” hypothesis is supported by the observation of higher populations of Phyllotreta spp., acting as facultative polyphages, in the Brassicaceae weed polyculture than in the non-Brassicaceae weed polyculture where Phyllotreta spp. are facultative monophages. Population densities of natural enemies (mostly coccinellids, carabids, and staphylinids) were higher in the polycultures than in the monoculture: carabid and staphylinid predators may be responsible for larval mortality in the imported cabbage worm, Pieris␣rapae, and in the diamondback larvae, Plutella xylostella. In spite of differences in densities of specialist herbivores across treatments, crop yield, leaf area (cm²), the proportion of leaf area damaged, and the number of leaves undamaged did not differ. These findings suggest that plant competition may interfere with attempts to reduce herbivore damage. We conclude that the use of weedy cultures can provide effective means of reducing herbivores if the crop and weed species are not related and plant competition is prevented. Received: 25 December 1995 / Accepted: 24 February 1997     

Complementarity in root architecture for nutrient uptake in ancient maize/bean and maize/bean/squash polycultures

Background and Aims

During their domestication, maize, bean and squash evolved in polycultures grown by small-scale farmers in the Americas. Polycultures often overyield on low-fertility soils, which are a primary production constraint in low-input agriculture. We hypothesized that root architectural differences among these crops causes niche complementarity and thereby greater nutrient acquisition than corresponding monocultures.

Methods

A functional–structural plant model, SimRoot, was used to simulate the first 40 d of growth of these crops in monoculture and polyculture and to determine the effects of root competition on nutrient uptake and biomass production of each plant on low-nitrogen, -phosphorus and -potassium soils.

Key Results

Squash, the earliest domesticated crop, was most sensitive to low soil fertility, while bean, the most recently domesticated crop, was least sensitive to low soil fertility. Nitrate uptake and biomass production were up to 7 % greater in the polycultures than in the monocultures, but only when root architecture was taken into account. Enhanced nitrogen capture in polycultures was independent of nitrogen fixation by bean. Root competition had negligible effects on phosphorus or potassium uptake or biomass production.

Conclusions

We conclude that spatial niche differentiation caused by differences in root architecture allows polycultures to overyield when plants are competing for mobile soil resources. However, direct competition for immobile resources might be negligible in agricultural systems. Interspecies root spacing may also be too large to allow maize to benefit from root exudates of bean or squash. Above-ground competition for light, however, may have strong feedbacks on root foraging for immobile nutrients, which may increase cereal growth more than it will decrease the growth of the other crops. We note that the order of domestication of crops correlates with increasing nutrient efficiency, rather than production potential.     

资源互补效应对多样性-生产力关系的影响

许多有关物种多样性-生态系统功能关系的观察、理论和实验研究都表明, 在局域尺度范围内, 植物种多样性对生态系统生产力存在正效应。 然而, 对于促成这种关系的潜在生态学机制却缺乏足够的了解。 该实验利用9种一年生栽培牧草, 采用各物种单播及混播的方法, 构建不同多样性梯度的实验群落, 对物种多样性与生态系统生产力的关系及资源互补效应对系统生产力的影响进行了研究。 结果表明, 在一年生植物群落内,植物种多样性在一定程度内对系统生产力存在正效应, 物种多样性与生产力呈二次函数关系, 关系式为y = -98.449x² + 1 039.2 x - 42.407, (R² = 0.423 1)。 各物种在资源利用、生长速度和竞争能力等功能特征方面存在较大差异, 最高产物种和最低产物种间产量相差5.8倍。 在同一多样性梯度内, 不同物种组合的群落间生产力和互补效应也存在较大差异, 说明物种的成分对生态系统生产力也有重要影响。 同时,在混播群落中程度不同地存在着资源的互补性利用, 说明物种多样性对系统生产力有增强作用, 但相关分析表明, 互补效应和物种多样性间不存在显著相关关系。互补效应的4种计算方法所反映的资源互补程度有所不同, 每种方法各有利弊, 在对系统的多样性效应作用机制进行评价时, 应根据具体情况, 同时采用几种方法, 以利于对资源互补效应做出恰当的估测。     

Herbivore response to vegetational diversity: spatial interaction of resources and natural enemies

Vegetational diversity in agricultural systems is predicted to reduce herbivore populations, but we observed the opposite effect: higher nymph population densities of a functionally monophagous herbivore, the squash bug, Anasa tristis (Hemiptera: Coreidae) in a vegetationally diverse squash-bean-corn polyculture than in a squash monoculture. We examined spatial and temporal aspects of squash bug and predator populations in relation to vegetational diversity. Average colonization, oviposition, and mortality rates for the herbivore were similar in monocultures and polycultures. In the polyculture, however, we found that squash bugs eggs were highly aggregated on plants on the outer edges of plots. Predation was also lower on plants near the edges, allowing the large aggregations of eggs found in the polyculture to escape predation and ultimately produce more squash bugs. Spatial interactions between herbivores and natural enemies may underlie some of the general effects of vegetational diversity on herbivores.     

Does diversity beget diversity? A case study of crops and weeds

Abstract. The ecological literature is ambiguous as to whether the initial diversity of a plant community facilitates or deters the diversity of colonizing species. We experimentally planted annual crop species in monoculture and polyculture, and examined the resulting weed communities. The species composition of weeds was similar among treatments, but the species richness of weeds was significantly higher in the polycultures than in the monocultures. This supports the ‘diversity begets diversity’ hypothesis. Environmental microheterogeneity, diversity promoters, and ecological equivalency do not seem able to explain the observed patterns.     

Nitrogen Fertilization Effects on Productivity and Nitrogen Loss in Three Grass-Based Perennial Bioenergy Cropping Systems

Nitrogen (N) fertilization can greatly improve plant productivity but needs to be carefully managed to avoid harmful environmental impacts. Nutrient management guidelines aimed at reducing harmful forms of N loss such as nitrous oxide (N₂O) emissions and nitrate (NO₃^-) leaching have been tailored for many cropping systems. The developing bioenergy industry is likely to make use of novel cropping systems, such as polycultures of perennial species, for which we have limited nutrient management experience. We studied how a switchgrass (Panicum virgatum) monoculture, a 5-species native grass mixture and an 18-species restored prairie responded to annual fertilizer applications of 56 kg N ha^-1 in a field-scale agronomic trial in south-central Wisconsin over a 2-year period. We observed greater fertilizer-induced N₂O emissions and sub-rooting zone NO₃^- concentrations in the switchgrass monoculture than in either polyculture. Fertilization increased aboveground net primary productivity in the polycultures, but not in the switchgrass monoculture. Switchgrass was generally more productive, while the two polycultures did not differ from each other in productivity or N loss. Our results highlight differences between polycultures and a switchgrass monoculture in responding to N fertilization.     

Does a multi‐plant diet benefit a polyphagous herbivore? A case study with Bemisia tabaci

Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) is a highly polyphagous herbivore. This research was conducted to compare the development of B. tabaci reared in a multi‐plant treatment (polyculture) with those in single‐plant treatments (monocultures). Adult B. tabaci females fed on a mixture of tomato, cabbage, cotton, cucumber, and kidney bean survived longer and laid more eggs than those fed exclusively on one of these plant species. Egg numbers per plant laid in the polyculture treatment were positively correlated with those laid on the same plant species in the monoculture treatments, and egg numbers per plant laid on tomato, cotton, and cucumber in the polyculture were significantly higher than those laid on the same plants in the monocultures. Concentrations of total protein and trehalose in B. tabaci were not significantly different after 7 days of feeding in the respective treatments, but activities of superoxide dismutases (SOD) and alkaline phosphatase (AKP) of B. tabaci in polyculture were lower than those in monoculture. Conversely, activities of trehalase, sucrase, and amylase in B. tabaci kept in polyculture were higher than those of insects from the monoculture. In each of the monoculture treatments, there was a negative correlation between AKP in B. tabaci and oviposition, and also between AKP and amylase. SOD and sucrase activities in B. tabaci were positively correlated with polyphenol oxidase (PPO) and peroxidase (POD) activities in plants. In the plants damaged by whiteflies in the polyculture treatment, activities of SOD in cucumber, PPO in cotton and kidney bean, and POD in tomato and cucumber were lower than those in the monoculture treatments, whereas SOD in cabbage and catalase (CAT) in tomato in the polyculture treatment were higher than those in the monoculture treatments.     

Productivity and complementarity in grassland microcosms of varying diversity

Several researchers have hypothesized that, through various mechanisms, loss of species and functional group richness from a plant community will affect the magnitude and interannual variability of productivity. To test this hypothesis, I conducted a microcosm study of California grassland communities that differed in species richness. I grew cohorts of microcosms that simulated undisturbed grassland (in one year) and gopher-disturbed grassland (in two consecutive years). As the number of species per functional group decreased from 4 to 1, biomass production remained constant in all three cohorts. As species richness decreased from 16 to 1 (or 8 to 1, in either case including a drop in functional group richness), productivity declined in one of the cohorts. In this cohort, productivity of one polyculture marginally exceeded that of the most productive monoculture. Resource complementarity and a type of selection effect may have each contributed to the observed diversity-productivity relationships. Results suggest the existence of a selection effect that involves species that are highly productive in mixtures, rather than in monoculture. Over two seasons, species and functional group richness did not affect the interannual variability of biomass production. Comparisons of interannual changes in the productivity of monocultures and polycultures suggested that, in some polycultures, increased water availability might have relieved interspecific competition more than intraspecific competition. Based on results from this experiment and other manipulative experiments, I develop a framework to explain the relationship between species richness and productivity in terrestrial plant communities. The framework highlights the importance of environmental variation in shaping the diversity/productivity relationship.     

Subinhibitory Antibiotic Concentrations Mediate Nutrient Use and Competition among Soil Streptomyces

Though traditionally perceived as weapons, antibiotics are also hypothesized to act as microbial signals in natural habitats. However, while subinhibitory concentrations of antibiotics (SICA) are known to shift bacterial gene expression, specific hypotheses as to how SICA influence the ecology of natural populations are scarce. We explored whether antibiotic ‘signals’, or SICA, have the potential to alter nutrient utilization, niche overlap, and competitive species interactions among Streptomyces populations in soil. For nine diverse Streptomyces isolates, we evaluated nutrient utilization patterns on 95 different nutrient sources in the presence and absence of subinhibitory concentrations of five antibiotics. There were significant changes in nutrient use among Streptomyces isolates, including both increases and decreases in the capacity to use individual nutrients in the presence vs. in the absence of SICA. Isolates varied in their responses to SICA and antibiotics varied in their effects on isolates. Furthermore, for some isolate-isolate-antibiotic combinations, competition-free growth (growth for an isolate on all nutrients that were not utilized by a competing isolate), was increased in the presence of SICA, reducing the potential fitness cost of nutrient competition among those competitors. This suggests that antibiotics may provide a mechanism for bacteria to actively minimize niche overlap among competitors in soil. Thus, in contrast to antagonistic coevolutionary dynamics, antibiotics as signals may mediate coevolutionary displacement among coexisting Streptomyces, thereby hindering the emergence of antibiotic resistant phenotypes. These results contribute to our broad understanding of the ecology and evolutionary biology of antibiotics and microbial signals in nature.     

Biodiversity improves the ecological design of sustainable biofuel systems

For algal biofuels to become a commercially viable and sustainable means of decreasing greenhouse gas emissions, growers are going to need to design feedstocks that achieve at least three characteristics simultaneously as follows: attain high yields; produce high quality biomass; and remain stable through time. These three qualities have proven difficult to achieve simultaneously under the ideal conditions of the laboratory, much less under field conditions (e.g., outdoor culture ponds) where feedstocks are exposed to highly variable conditions and the crop is vulnerable to invasive species, disease, and grazers. Here, we show that principles from ecology can be used to improve the design of feedstocks and to optimize their potential for “multifunctionality.” We performed a replicated experiment to test these predictions under outdoor conditions. Using 80 ponds of 1,100 L each, we tested the hypotheses that polycultures would outperform monocultures in terms of the following functions: biomass production, yield of biocrude from biomass, temporal stability, resisting population crashes, and resisting invasions by unwanted species. Overall, species richness improved stability, biocrude yield, and resistance to invasion. While this suggests that polycultures could outperform monocultures on average, invasion resistance was the only function where polycultures outperformed the best single species in the experiment. Due to tradeoffs among different functions that we measured, no species or polyculture was able to maximize all functions simultaneously. However, diversity did enhance the potential for multifunctionality—the most diverse polyculture performed more functions at higher levels than could any of the monocultures. These results are a key finding for ecological design of sustainable biofuel systems because they show that while a monoculture may be the optimal choice for maximizing short‐term biomass production, polycultures can offer a more stable crop of the desired species over longer periods of time.     

Overyielding in grassland communities: testing the sampling effect hypothesis with replicated biodiversity experiments

We derive and test some assumptions and predictions of the Sampling Effect Hypothesis (SEH) by examining the relationship between the traits of species in monoculture and their relative abundance in mixture, and by comparing polyculture performance with single-species plots. Although we found a positive relationship between production in monoculture and dominance in mixtures as predicted by the SEH, the relationship had low explanatory power. Counter to predictions, the species with the highest monoculture biomass were not able to strongly dominate all mixtures; instead the dominance of these species decreased with increasing species richness. On average, polycultures did not achieve greater biomass than (transgressively overyield) the species in each mixture, or at each site, that was most productive in monoculture. However, mixture yields did transgressively overyield both the monoculture biomass of the dominant species in the mixtures, and the weighted average of all monocultures (non-transgressive overyielding), both of which were positively related to increasing species richness. The varying responses of different overyielding tests resulted because resource partitioning and positive interactions were often counter-balanced by selection for species with lower biomass than the highest-yielding monocultures. Judging whether or not mixtures overyield therefore depends in part upon which species is the basis for comparison. We present a new general framework for overyielding analysis where every monoculture provides a potential comparison and from which the most relevant tests can be selected.     

The long-term relationship between plant diversity and total plant biomass depends on the mechanism maintaining diversity

The relationship between plant diversity and total plant biomass is of great current interest in ecology. Niche differences among plants are widely thought to promote both plant coexistence, and higher biomass in more diverse polycultures. Using simple mechanistic models, I demonstrate that not all niche differences among plants are equally likely to promote high total biomass in polyculture. In particular, transgressive overyielding (when a polyculture of plants outperforms any monoculture) occurs for a wide range of parameter values when plants coexist in polyculture due to differences in resource use. Transgressive overyielding occurs only for a limited set of parameter values when specialist herbivores mediate plant coexistence, and is impossible when generalist herbivores mediate coexistence. Niche differences among plants promote high biomass in polyculture only when plants coexist in polyculture at the expense of other trophic levels – that is, by converting into biomass resources that would otherwise be bound in herbivores, or exist in a free state. A major challenge for future work will be to identify the joint consequences of different coexistence mechanisms for plant diversity and ecosystem performance.     

Soil microbial diversity and soil functioning affect competition among grasses in experimental microcosms

A gradient of microbial diversity in soil was established by inoculating pasteurized soil with microbial populations of different complexity, which were obtained by a combination of soil fumigation and filtering techniques. Four different soil diversity treatments were planted with six different grass species either in monoculture or in polyculture to test how changes of general microbial functions, such as catabolic diversity and nutrient recycling efficiency would affect the performance of the plant communities. Relatively harsh soil treatments were necessary to elicit visible effects on major soil processes such as decomposition and nitrogen cycling due to the high redundancy and resilience of soil microbial communities. The strongest effects of soil diversity manipulations on plant growth occurred in polycultures where interspecific competition between plants was high. In polycultures, soil diversity reduction led to a gradual, linear decline in biomass production of one subordinate grass species (Bromus hordeaceus), which was compensated by increased growth of two intermediate competitors (Aegilops geniculata, B. madritensis). This negative covariance in growth of competing grass species smoothed the effects of soil diversity manipulations at the plant community level. As a result, total shoot biomass production remained constant. Apparently the effects of soil diversity manipulations were buffered because functional redundancy at both, the microbial and the plant community level complemented each other. The results further suggests that small trade-offs in plant fitness due to general functional shifts at the microbial level can be significant for the outcome of competition in plant communities and thus diversity at much larger scales.     

Associational plant refuges: convergent patterns in marine and terrestrial communities result from differing mechanisms

Summary An associational plant refuge occurs when a plant that is susceptible to herbivory gains protection from herbivory when it is associated with another plant. In coastal North Carolina, the abundance of the palatable red alga Gracilaria tikvahiae is positively correlated with the abundance of the unpalatable brown alga Sargassum filipendula during times of increased herbivore activity. To see if grazing by the sea urchin Arbacia punctulata could generate this pattern, controlled experiments were conducted in out-door microcosms and in the laboratory. Gracilaria beneath a canopy of Sargassum was eaten significantly less than Gracilaria alone. When Arbacia were excluded, Gracilaria alone grew significantly more than Gracilaria beneath Sargassum, demonstrating that Sargassum is a competitor of Gracilaria. Experiments investigating Sargassum's deterrent role indicated that Sargassum decreased the foraging range of Arbacia and the rate at which it fed on Gracilaria. Additional experiments with plastic Sargassum mimics indicated that the decreased grazing on Gracilaria was not a result of Sargassum morphology, but was probably attributable to some chemical characteristic of Sargassum. The pattern of increased grazing in monocultures (only Gracilaria present) versus polycultures (both Gracilaria and Sargassum present) demonstrated in this study also has been demonstrated for plant-insect interactions in terrestrial communities. In these communities, insect density is higher in monocultures than in polycultures because insects find and immigrate to monocultures more rapidly, and once in a monoculture, they emigrate from them less often than from polycultures. In this study, urchins did not find and immigrate to monocultures more rapidly, nor did they tend to stay in them once they were found; in fact, they emigrated from monocultures of Gracilaria more rapidly than from Gracilaria and Sargassum polycultures. Increased grazing in Gracilaria monocultures resulted from increased rates of movement and feeding of individual herbivores, not from increased herbivore density as has been reported for terrestrial systems.     

Effects of plant diversity and time of colonization on an herbivore-plant interaction

Summary Experimental field plantings showed that plant diversity strongly affected the population dynamics of a specialist herbivore, the striped cucumber beetle, Acalymma vittata (Fab.) (Coleoptera: Chrysomelidae). Population densities over time were characterized by two peaks in numbers (from colonization and reproduction, respectively) and were consistently higher in cucumber monocultures (Cucumis sativus L.) than in polycultures of cucumbers, corn (Zea mays L.), and broccoli (Brassica oleracea L.). Greater abundances in monocultures appear to result from two factors: (1) per individual reproductive rates were greater in monocultures than in polycultures, and (2) mark-recapture studies confirmed that beetles stay in monocultures for a longer period of time than in polycultures. Differences in predation did not appear to contribute to the overall differences in herbivore abundances.The primary impact of A. vittata on its host plant, C. sativus, is the dissemination of bacterial wilt disease, Erwinia tracheiphila (E.F.Sm.). Greater numbers of beetles led to greater plant mortality in monocultures. It is suggested that factors other than numbers of beetles (e.g., shading, allelopathy, microclimate) are more important in influencing plant reproduction, since cucumber plants in monocultures had greater yields than did plants grown in polycultures. However, time of beetle colonization strongly affected plant parameters, indicating that the length of time during which herbivores are interacting with plants is of critical importance to plant survivorship, and thus reproduction.     

Temporal variability in nutrient use among Streptomyces suggests dynamic niche partitioning

Soil bacteria spend significant periods in dormant or semi-dormant states that are interrupted by resource pulses which can lead to periods of rapid growth and intense nutrient competition. Microbial populations have evolved diverse strategies to circumvent competitive interactions and facilitate coexistence. Here, we show that nutrient use of soilborne Streptomyces is temporally partitioned during experimental resource pulses, leading to reduced niche overlap, and potential coexistence. Streptomyces grew rapidly on the majority of distinct 95 carbon sources but varied in which individual resources were utilized in the first 24 h. Only a handful of carbon sources (19 out of 95) were consistently utilized (>95% of isolates) most rapidly in the first 24 h. These consistently utilized carbon sources also generated the majority of biomass accumulated by isolates. Our results shed new light on a novel mechanism microbes may employ to alleviate competitive interactions by temporally partitioning the consumption of carbon resources. As competitive interactions have been proposed to drive the suppression of disease-causing microbes in agronomic soils, our findings may hold widespread implications for soil management for plant health.     

Density of Anomoea flavokansiensis on Desmanthus illinoensis in monoculture and polyculture

The effect of plant species diversity on the density of the herbivore, Anomoea flavokansiensis Moldenke (Coleoptera: Chrysomelidae), on Desmanthus illinoensis (Michaux) MacMillan (Mimosaceae), a promising North American legume for exploring the principles of diverse, perennial grain agriculture was examined. From mid-June to early August A. flavokansiensis feeds on young leaves and inflorescences of D. llinoensis. At high density, A. flavokansiensis potentially reduces seed yield and is thus an important consideration for long-term stands that are to be grown without insecticides. The potential to manage this insect via intercropping its host species with other, non-host perennial species by monitoring A. flavokansiensis density on D. illinoensis within experimental monocultures, two-species mixtures with Tripsacum dactyloides, and three-species mixtures with T. dactyloides and Leymus racemosus was examined. Insects were censused 2–3 times weekly from mid-June to early August at two sites from 1991 to 1995. In the first three years, beetle density was generally low (<1 per plant), and did not differ among treatments. In the fourth year, however, beetle density peaked at 15 and 25 insects per plant at the two sites, and was highest within monoculture for most dates. In 1995, density was again low, but tended to remain higher in monoculture at one site. The results suggest that beetle density on D. illinoensis can be reduced in polyculture and may hold promise for the management of this insect herbivore within perennial grain polycultures.     

Effects of polyculture and monoculture farming in oil palm smallholdings on tropical fruit‐feeding butterfly diversity

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Inhibitory and nutrient use phenotypes among coexisting Fusarium and Streptomyces populations suggest local coevolutionary interactions in soil

Bacteria and fungi are key components of virtually all natural habitats, yet the significance of fungal-bacterial inhibitory interactions for the ecological and evolutionary dynamics of specific bacterial and fungal populations in natural habitats have been overlooked. More specifically, despite the broad consensus that antibiotics play a key role in providing a fitness advantage to competing microbes, the significance of antibiotic production in mediating cross-kingdom coevolutionary interactions has received relatively little attention. Here, we characterize reciprocal inhibition among Streptomyces and Fusarium populations from prairie soil, and explore antibiotic inhibition in relation to niche overlap among sympatric and allopatric populations. We found evidence for local adaptation between Fusarium and Streptomyces populations as indicated by significantly greater inhibition among sympatric than allopatric populations. Additionally, for both taxa, there was a significant positive correlation between the strength of inhibition against the other taxon and the intensity of resource competition from that taxon among sympatric but not allopatric populations. These data suggest that coevolutionary antagonistic interactions between Fusarium and Streptomyces are driven by resource competition, and support the hypothesis that antibiotics act as weapons in mediating bacterial–fungal interactions in soil.     

Bioconversion of filter mud using vermicomposting employing two exotic and one local earthworm species

Three different earthworm species Eisenia fetida, Eudrilus eugeniae and Perionyx excavatus in individual (Monocultures) and combinations (Polycultures) were utilized to compare the suitability of worm species for vermicomposting of filter mud as well as the quality of the end product. The filter mud blended with saw dust can be directly converted into good quality fertilizer (vermicompost). Eight different reactors including three monocultures and four polycultures of E. fetida, E. eugeniae and P. excavatus and one control were used for the experiment. Vermicomposting resulted in significant reduction in C/N ratio, pH, total organic matter (TOC) but increase in electrical conductivity (EC), total nitrogen (TN), total phosphorus (TP) and macronutrients (K, Ca and Na). Oxygen uptake rate (OUR) dropped up to 1.64–1.95 mg/g (volatile solids) VS/day for monoculture reactors and 1.45–1.78 mg/g VS/day for polycultures reactors, respectively, after 45 days of vermicomposting. Cocoon production and the earthworm biomass increased as vermicomposting progressed. On an overall the mono as well as polyculture reactors produced high quality stable compost free from pathogens and no specific differentiation could be inferred between the reactors.