Bacterivores, mostly represented by protists and nematodes, are a key component of soil biodiversity involved in soil fertility and plant productivity. In the current context of global change and soil biodiversity erosion, it becomes urgent to suitably recognize and quantify their ecological importance in ecosystem functioning. Using meta-analysis tools, we aimed at providing a quantitative synthesis of the ecological importance of soil bacterivores on ecosystem functions. We also intended to produce an overview of the ecological factors that are expected to drive the magnitude of bacterivore effects on ecosystem functions. Bacterivores in soil contributed significantly to numerous key ecosystem functions. We propose a new theoretical framework based on ecological stoichiometry stressing the role of C:N:P ratios in soil, microbial and plant biomass as important parameters driving bacterivore-effects on soil N and P availability for plants, immobilization of N and P in the bacterial biomass, and plant responses in nutrition and growth.

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Ecological importance of soil bacterivores for ecosystem functions

For decades, the term “rhizosphere fauna” has been used as a synonym to denote agricultural pests among root herbivores, mainly nematodes and insect larvae. We want to break with this constrictive view, since the connection between plants and rhizosphere fauna is far more complex than simply that of resource and consumer. For example, plant roots have been shown to be neither defenceless victims of root feeders, nor passive recipients of nutrients, but instead play a much more active role in defending themselves and in attracting beneficial soil microorganisms and soil fauna. Most importantly, significant indirect feed-backs exist between consumers of rhizosphere microorganisms and plant roots. In fact, the majority of soil invertebrates have been shown to rely profoundly on the carbon inputs from roots, breaking with the dogma of soil food webs being mainly fueled by plant litter input from aboveground. In this review we will highlight areas of recent exciting progress and point out the black boxes that still need to be illuminated by rhizosphere zoologists and ecologists.

Rhizosphere fauna: the functional and structural diversity of intimate interactions of soil fauna with plant roots

In the context of increasing global food demand, ecological intensification of agroecosystems is required to increase nutrient use efficiency in plants while decreasing fertilizer inputs. Better exploration and exploitation of soil resources is a major issue for phosphorus, given that rock phosphate ores are finite resources, which are going to be exhausted in decades from now on. We review the processes governing the acquisition by plants of poorly mobile nutrients in soils, with a particular focus on processes at the root–soil interface. Rhizosphere processes are poorly accounted for in most plant nutrition models. This lack largely explains why present-day models fail at predicting the actual uptake of poorly mobile nutrients such as phosphorus under low input conditions. A first section is dedicated to biophysical processes and the spatial/temporal development of the rhizosphere. A second section concentrates on biochemical/biogeochemical processes, while a third section addresses biological/ecological processes operating in the rhizosphere. New routes for improving soil nutrient efficiency are addressed, with a particular focus on breeding and ecological engineering options. Better mimicking natural ecosystems and exploiting plant diversity appears as an appealing way forward, on this long and winding road towards ecological intensification of agroecosystems.

Acquisition of phosphorus and other poorly mobile nutrients by roots. Where do plant nutrition models fail

It is increasingly believed that substantial soil organic carbon (SOC) can be sequestered in conservation tillage system by manipulating the functional groups of soil biota. Soil aggregates of different size provide diverse microhabitats for soil biota and consequently influence C sequestration. Our objective was to evaluate the contributions of soil biota induced by tillage systems to C sequestration among different aggregate size fractions. Soil microbial and nematode communities were examined within four aggregate fractions: large macroaggregates (>2 mm), macroaggregates (2e1 mm), small macroaggregates (1e0.25 mm) and microaggregates ( 1 mm aggregate fractions were different from those in 1m m aggregates, while more gram-positive bacteria and plant-parasitic nematodes might increase C accumulation within <1 mm aggregates. Our findings suggested that the increase in microbial biomass and nematode abundance and the alteration in their community composition at the micro-niche within aggregates could contribute to the higher C sequestration in conservation tillage systems (NT and RT). 2013 Elsevier Ltd. All rights reserved.

Contributions of soil biota to C sequestration varied with aggregate fractions under different tillage systems

Effects of organic amendment and tillage on soil microorganisms and microfauna

Bacterial-feeding nematodes constitute one of the primary grazers of soil bacteria. We investigated the effects of selective grazing of a representative nematode (Zeldia punctata, Cephalobidae) on nematode life history and population biology and on the soil microbial community. Firstly, we measured (i) the effect of five different bacterial strains on the nematode life cycle using petri dishes and (ii) the impact of bacterial inoculation on nematode population growth in a soil microcosm. Selection of the five bacterial strains was based on morphology, cell-wall characteristics and mucus production. Z. punctata development was strongly affected by the type of bacteria ingested, independent of experimental design. Bacterial cell-wall characteristics seemed to directly affect Z. punctata development since high nematode densities were only reached with gram-negative strains (Pseudomonas monteilii and Methylobacterium nodulans). In petri dishes, the filamentous organisms (Actinomyces sp.) and mucus-producing bacteria (Bradyrhizobium sp.) led to the least reproduction. Duration of the various nematode life phases (egg, juvenile, reproductive stage and non-reproductive stage) was significantly affected by the bacterial food source. Total life span varied from 12.5 days (Bradyrhizobium sp.) to 40 days (Pseudomonas monteilii). Secondly, we monitored the influence of Z. punctata on the indigenous soil microbial community in the presence or absence of a maize plantlet. Nematode inoculation led to an increase in bacterial activity (as measured by alkaline phosphatase activity) but did not significantly influence bacterial biomass. The genetic fingerprint (DGGE) of soil bacteria was more influenced by plant presence than by nematode inoculation. Nematode activity has important repercussions on N flux in the soil since inoculation of Z. punctata in the absence of plants resulted in a net increase of N mineralization (2 mg N per pot) while a decrease of mineral N (0.5 mg N per pot) was observed in the absence of the nematodes, due to bacterial immobilization. This study underscores the close relationship between selective bacterial grazing and nematode development. Nevertheless, the impact of nematode grazing on the overall soil microbial community seems to primarily affect microbial activity and relative dominance rather than microbial diversity.

Interactions between Zeldia punctata (Cephalobidae) and bacteria in the presence or absence of maize plants

Nutrition on bacteria by bacterial-feeding nematodes and consequences on the structure of soil bacterial community

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Interactions between Zeldia punctata (Cephalobidae) and bacteria in the presence or absence of maize plants

Nutrition on bacteria by bacterial-feeding nematodes and consequences on the structure of soil bacterial community