Priming effects are strong short-term changes in the turnover of soil organic matter caused by comparatively moderate treatments of the soil. In the course of priming effects large amounts of C, N and other nutrients can be released or immobilized in soil in a very short time. These effects have been measured in many field and laboratory experiments; however, only a few of the studies were aimed at an extended investigation of the mechanisms of such phenomena. The aim of this overview is to reveal possible causes and processes leading to priming actions using the references on agricultural ecosystems and model experiments. Multiple mechanisms and sources of released C and N are presented and summarized in Tables for positive and negative real and apparent priming effects induced after the addition of different organic and mineral substances to the soil. Soil microbial biomass plays the key role in the processes leading to the real priming effects. The most important mechanisms for the real priming effects are the acceleration or retardation of soil organic matter turnover due to increased activity or amount of microbial biomass. Isotopic exchange, pool substitution, and different uncontrolled losses of mineralized N from the soil are responsible for the apparent N priming effects. Other multiple mechanisms (predation, competition for nutrients between roots and microorganisms, preferred uptake, inhibition, etc.) in response to addition of different substances are also discussed. These mechanisms can be distinguished from each other by the simultaneous monitoring of C and N release dynamics; its comparison with the course of microbial activity; and by the labelling of different pools with 14 C or 13 C and 15 N. Quantitative methods for describing priming effects and their dynamics using 14 C and 15 N isotopes, as well as for non-isotopic studies are proposed.

Review of mechanisms and quantification of priming effects.

In this re-evaluation of our 10-year old paper on priming effects, I have considered the latest studies and tried to identify the most important needs for future research. Recent publications have shown that the increase or decrease in soil organic matter mineralization (measured as changes of CO 2 efflux and N mineralization) actually results from interactions between living (microbial biomass) and dead organic matter. The priming effect (PE) is not an artifact of incubation studies, as sometimes supposed, but is a natural process sequence in the rhizosphere and detritusphere that is induced by pulses or continuous inputs of fresh organics. The intensity of turnover processes in such hotspots is at least one order of magnitude higher than in the bulk soil. Various prerequisites for high-quality, informative PE studies are outlined: calculating the budget of labeled and total C; investigating the dynamics of released CO 2 and its sources; linking C and N dynamics with microbial biomass changes and enzyme activities; evaluating apparent and real PEs; and assessing PE sources as related to soil organic matter stabilization mechanisms. Different approaches for identifying priming, based on the assessment of more than two C sources in CO 2 and microbial biomass, are proposed and methodological and statistical uncertainties in PE estimation and approaches to eliminating them are discussed. Future studies should evaluate directions and magnitude of PEs according to expected climate and land-use changes and the increased rhizodeposition under elevated CO 2 as well as clarifying the ecological significance of PEs in natural and agricultural ecosystems. The conclusion is that PEs – the interactions between living and dead organic matter – should be incorporated in models of C and N dynamics, and that microbial biomass should regarded not only as a C pool but also as an active driver of C and N turnover.

Priming effects : interactions between living and dead organic matter

It is generally accepted that the low quality of soil carbon limits the amount of energy available for soil microorganisms, and in turn the rate of soil carbon mineralization. The priming effect, i.e. the increase in soil organic matter (SOM) decomposition rate after fresh organic matter input to soil, is often supposed to result from a global increase in microbial activity due to the higher availability of energy released from the decomposition of fresh organic matter. Work to date, however, suggests that supply of available energy induces no effect on SOM mineralization. The mechanisms of the priming effect are much more complex than commonly believed. The objective of this review was to build a conceptual model of the priming effect based on the contradictory results available in the literature adopting the concept of nutritional competition. After fresh organic matter input to soils, many specialized microorganisms grow quickly and only decompose the fresh organic matter. We postulated that the priming effect results from the competition for energy and nutrient acquisition between the microorganisms specialized in the decomposition of fresh organic matter and those feeding on polymerised SOM.

The priming effect of organic matter: a question of microbial competition

Traditional models of soil organic matter (SOM) decomposition are all based on first order kinetics in which the decomposition rate of a particular C pool is proportional to the size of the pool and a simple decomposition constant (dC/dt=kC). In fact, SOM decomposition is catalyzed by extracellular enzymes that are produced by microorganisms. We built a simple theoretical model to explore the behavior of the decomposition–microbial growth system when the fundamental kinetic assumption is changed from first order kinetics to exoenzymes catalyzed decomposition (dC/dt=KC×Enzymes). An analysis of the enzyme kinetics showed that there must be some mechanism to produce a non-linear response of decomposition rates to enzyme concentration—the most likely is competition for enzyme binding on solid substrates as predicted by Langmuir adsorption isotherm theory. This non-linearity also induces C limitation, regardless of the potential supply of C. The linked C and N version of the model showed that actual polymer breakdown and microbial use of the released monomers can be disconnected, and that it requires relatively little N to maintain the maximal rate of decomposition, regardless of the microbial biomass’ ability to use the breakdown products. In this model, adding a pulse of C to an N limited system increases respiration, while adding N actually decreases respiration (as C is redirected from waste respiration to microbial growth). For many years, researchers have argued that the lack of a respiratory response by soil microbes to added N indicates that they are not N limited. This model suggests that conclusion may be wrong. While total C flow may be limited by the functioning of the exoenzyme system, actual microbial growth may be N limited.

The implications of exoenzyme activity on microbial carbon and nitrogen limitation in soil: a theoretical model

The knowns, known unknowns and unknowns of sequestration of soil organic carbon

Glucose, alfalfa insolubles, and complete alfalfa uniformly labeled with carbon-14 were added to an organic soil to determine under controlled conditions, if the loss of the soil due to biological decomposition could be counteracted. Carbon-14 labeled material was used so that it could be determined whether or not there was an increase in the decomposition of the native organic matter as a result of the added material, and, if this was true, whether the soluble fraction of the added materials, or the insoluble fraction, was chiefly responsible. The materials were added, the soil incubated, and CO₂ collected at intervals. The insoluble fraction of alfalfa caused a larger breakdown of native organic matter than solubles alone or complete alfalfa. It was impossible to build up the organic content of the soil, under these conditions, but the net loss of carbon was diminished by the addition of the organic materials.

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The Effect of the Addition of Organic Materials on the Decomposition of an Organic Soil

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The Effect of the Addition of Organic Materials on the Decomposition of an Organic Soil