Five main biogenic sources of CO2 efflux from soils have been distinguished and described according to their turnover rates and the mean residence time of carbon. They are root respiration, rhizomicrobial respiration, decomposition of plant residues, the priming effect induced by root exudation or by addition of plant residues, and basal respiration by microbial decomposition of soil organic matter (SOM). These sources can be grouped in several combinations to summarize CO2 efflux from the soil including: root-derived CO2, plant-derived CO2, SOM-derived CO2, rhizosphere respiration, heterotrophic microbial respiration (respiration by heterotrophs), and respiration by autotrophs. These distinctions are important because without separation of SOM-derived CO2 from plant-derived CO2, measurements of total soil respiration have very limited value for evaluation of the soil as a source or sink of atmospheric CO2 and for interpreting the sources of CO2 and the fate of carbon within soils and ecosystems. Additionally, the processes linked to the five sources of CO2 efflux from soil have various responses to environmental variables and consequently to global warming. This review describes the basic principles and assumptions of the following methods which allow SOMderived and root-derived CO2 efflux to be separated under laboratory and field conditions: root exclusion techniques, shading and clipping, tree girdling, regression, component integration, excised roots and in situ root respiration; continuous and pulse labeling, 13 C natural abundance and FACE, and radiocarbon dating and bomb- 14 C. A short sections cover the separation of the respiration of autotrophs and that of heterotrophs, i.e. the separation of actual root respiration from microbial respiration, as well as methods allowing the amount of CO2 evolved by decomposition of plant residues and by priming effects to be estimated. All these methods have been evaluated according to their inherent disturbance of the ecosystem and C fluxes, and their versatility under various conditions. The shortfalls of existing approaches and the need for further development and standardization of methods are highlighted. q 2005 Elsevier Ltd. All rights reserved.

Sources of CO2 efflux from soil and review of partitioning methods

Modification of Ecosystems by Ungulates

Understanding how terrestrial ecosystems function requires a combined aboveground–belowground approach, because of the importance of feedbacks that occur between herbivores, producers, and the decomposer subsystem. In this paper, we identify several mechanisms by which herbivores can indirectly affect decomposer organisms and soil processes through altering the quantity and quality of resources entering the soil. We show that these mechanisms are broadly similar in nature for both foliar and root herbivory, regardless of whether they operate in the short term as a result of physiological responses of individual plants to herbivore attack or long-term following alteration of plant community structure by herbivores and subsequent changes in the quality of litter inputs to soil. We propose that a variety of possible mechanisms is responsible for the idiosyncratic nature of herbivore effects on soil biota and ecosystem function; positive, negative, or neutral effects of herbivory are possible depending upon the balance of these different mechanisms. However, we predict that positive effects of herbivory on soil biota and soil processes are most common in ecosystems of high soil fertility and high consumption rates, whereas negative effects are most common in unproductive ecosystems with low consumption rates. The significance of multiple-species herbivore communities is also emphasized, and we propose that if resource use complementarity among herbivore species or functional groups leads to greater total consumption of phytomass, and thus greater net herbivory, then both positive and negative consequences of increasing herbivore diversity for belowground properties and processes are theoretically possible. Research priorities are highlighted and include a need for comparative studies of herbivore impacts on above- and belowground processes across ecosystems of varying productivity, as well as a need for experimental testing of the influence of antiherbivore defense compounds on complex multitrophic interactions in the rhizosphere and the significance of multiple herbivore species communities on these plant–soil interactions.

Herbivore-mediated linkages between aboveground and belowground communities

Grazing can alter the spatial heterogeneity of vegetation, influencing ecosystem processes and biodiversity. Our objective was to identify why grazing causes increases in the spatial heterogeneity of vegetation in some cases, but decreases in others. The immediate effect of grazing on heterogeneity depends on the interaction between the spatial pattern of grazing and the pre-existing spatial pattern of vegetation. Depending on the scale of observation and on the factors that determine animal distribution, grazing patterns may be stronger or weaker than vegetation patterns, or may mirror the spatial structure of vegetation. For each possible interaction between these patterns, we make a prediction about resulting changes in the spatial heterogeneity of vegetation. Case studies from the literature support our predictions, although ecosystems characterized by strong plant-soil interactions present important exceptions. While the processes by which grazing causes increases in heterogeneity are clear, how grazing leads to decreases in heterogeneity is less so. To explore how grazing can consistently dampen the fine-scale spatial patterns of competing plant species, we built a cell-based simulation model that features two competing plant species, different grazing patterns, and different sources of vegetation pattern. Only the simulations that included neighborhood interactions as a source of vegetation pattern produced results consistent with the predictions we derived from the literature review.

The effect of grazing on the spatial heterogeneity of vegetation

Summary

1
Shrub encroachment has been widely observed in savanna regions. This study analysed the causes of shrub encroachment in the lowveld savanna of north-eastern Swaziland, southern Africa, and highlighted management regimes that can be used to reduce or prevent it.

2
The rates and dynamics of shrub encroachment were quantified for the period 1947–90 using aerial photographs, and for 1997 using a ground survey. Five similar areas with different land-use histories were compared to investigate the relative importance of fire, herbivory, rainfall, soil type and shrub density in driving shrub dynamics.

3
In the study area as a whole, shrub cover increased from a mean of 2% in 1947 to 31% in 1990. Dichrostachys cinerea accounted for most of the increase in cover, contributing 81% to total shrub cover during 1997. Shrub cover was strongly correlated with shrub density and weakly negatively correlated with tree cover.

4
Shrub encroachment varied across land-use fence lines. The key determinants of shrub dynamics were grazing, through its negative effect on fire frequency and an interaction between drought frequency and high shrub cover. Browsing pressure had a significant but minor impact on dynamics, while soil type had no significant effect. High grazing pressures through their effect on fire frequency were critical throughout the study period in promoting shrub encroachment, while the interaction between drought and high shrub cover produced declines in the later stages. Browsing had an impact on encroachment only in the early stages.

5
Frequent fires, facilitated by low grazing pressures, were capable of preventing shrub encroachment. When coupled with drought, frequent fires could reduce high shrub densities.

6
As cover and density were strongly correlated, it can be inferred from the negative correlation between change in cover (density) and initial cover (density) that the rate of shrub encroachment was cover (density)-dependent and that there was a shrub equilibrium of 40% cover, approximating to 2400 plants ha−1. Shrub population growth was driven by events (fire, drought) as well as continuous agents (density dependence, mean browsing and grazing pressure).

7
Bush encroachment can be reversed by a combination of management (frequent fires) and climatic events (drought). The implications for savanna management are discussed.

https://besjournals.onlinelibrary.wiley.com/doi/pdf/10.1046/j.1365-2664.2001.00567.x

Dynamics of shrub encroachment in an African savanna: relative influences of fire, herbivory, rainfall and density dependence

Plant responses to herbivory and links to belowground nitrogen cycling were investigated at Wind Cave National Park, South Dakota. Laboratory estimates of net nitrogen mineralization were highest in soils from the more altered areas of prairie dog colonies (Cynomys ludovicianus) and lowest in the adjacent, lightly grazed, uncolonized grassland. The ratio of CO2: net nitrogen mineralized, an index of immobilization, was highest in the uncolonized grassland and lowest in the altered core areas. Soil moisture was an important modifier of in situ field estimates of net nitrogen mineralization. Root biomass, an important carbon source for decomposers in perennial grasslands, was lowest in the altered core area and highest in the adjacent uncolonized grassland. Decreased nitrogen immobilization and increased net nitrogen mineralization in the laboratory in- cubations likely resulted from decreased root carbon inputs in grazed areas, which limited carbon availability to decomposers. Such increases in plant-available nitrogen may partially explain the frequently reported grazing-induced increases in shoot nitrogen concentrations. These studies suggest that carbon allocation to roots is a key link determining nitrogen- cycling responses to herbivory.

Elizabeth A. Holland

Papers

Plant Response to Herbivory and Belowground Nitrogen Cycling