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International Journal of Biodiversity Science, Ecosystem Services & Management, 2014 Vol. 10, No. 3, 177–186, http://dx.doi.org/10.1080/21513732.2014.926990 Soil-based ecosystem services: a synthesis of nutrient cycling and carbon sequestration assessment methods a,b a c Bhim B. Ghaley *, John R. Porter and Harpinder S. Sandhu Faculty of Science, Department of Plant and Environmental Sciences, University of Copenhagen, Højbakkegård Allé 30, 2630 Taastrup, Denmark; Copenhagen Plant Science Centre, Faculty of Science, University of Copenhagen, Thorvaldsensvej 40, 1871 Frederiksberg C, Denmark; School of the Environment, Flinders University, GPO Box 2100, Adelaide, SA 5001, Australia Among the soil-based ecosystem services (ES), nutrient cycling and carbon sequestration have direct influence on the biogeochemical cycles and greenhouse gas emissions affecting provision of other ES that support human existence. We reviewed methods to assess the two key ES by identifying their strengths and weaknesses and have made suggestions for using appropriate methods for better understanding of the ecosystem functions for the provision of ES. Relevant papers for the review were chosen on the basis of (i) diversity of studies on the two key ES in different ecosystems, (ii) methodologies applied and (iii) detailed descriptions of the trial locations in terms of vegetation, soil type, location and climatic information. We concluded that (i) elemental stoichiometrical ratios could be a potential approach to assess the health of ecosystems in terms of provision of the two ES discussed, (ii) stoichiometric imbalances need to be avoided between the supply and the demand of the nutrients to maintain the ES provision in terrestrial ecosystems and (iii) stoichiometric ratios can act as a management tool at a field, farm and at landscape level, to complement other compositional biodiversity and functional diversity approaches to ensure sustainable provision of ES. Keywords: ecosystem functions; litter decomposition; mineralisation; assessment methodologies; stoichiometry Introduction conducted in diverse ecosystems at different spatial and temporal scales using different methods (Costanza et al. Ecosystem functions deliver goods and services (ecosys- 1997; Sandhu et al. 2008, 2010a; Porter et al. 2009; tem services (ES)) that satisfy human needs, directly or Ghaley et al. 2014). Among the ES, nutrient cycling and indirectly (De Groot et al. 2002; Sandhu & Wratten 2013). carbon sequestration have direct influence on the biogeo- ES are the basis of economic and social well-being of chemical cycles (elemental/ stoichiometric ratios) and human beings (UNEP 2010; TEEB 2010). ES are classi- greenhouse gas (GHG) emission affecting provision of fied into four major categories (Table 1): (a) provisioning other ES (Ballantyne et al. 2008; Allen & Gillooly 2009). services like food, fibre, bio-energy, etc.; (b) supporting Several studies have investigated nutrient cycling in differ- services like soil fertility, nutrient cycling, provision of ent ecosystems like forests (Rożen et al. 2010), arable water, etc.; (c) regulating services like pollination, biolo- agriculture (Sandhu et al. 2008, 2010a) and permanent gical control of pests and diseases, etc.; and (d) cultural grassland (Hamel et al. 2007). The nutrient cycling ES is services like recreation, education (MEA 2005). ES provi- based on determination of major ecosystem functions like sion occurs at multiple scales from carbon sequestration nitrogen fixation, phosphorus supply by mycorrhizal fungi for climate regulation at the global scale to soil structure and litter decomposition and mineralisation rate. With maintenance and fertility at the local scale (MEA 2005). regard to nutrient cycling ES, litter decomposition and Due to the spatial and temporal nature of ES, multiple mineralisation is the most common ecosystem functions/ approaches are preferred over a single method for objective processes in terms of number of investigations (de Bello assessment of ES. Different ES are the end products of et al. 2010) and methods of assessment applied. Similarly, ecosystem functions, some of which can be estimated by carbon sequestration is also widely investigated ES due to on-site measurements in the field, whereas other ES of its role in GHG emission and climate change effects and social value can be better estimated by questionnaire inter- multitude of methods are used for its assessment. Given the view or survey methods. For example, provisioning, regu- multiple methods applied for a single ecosystem function lating and supporting ES like food, fodder and bio-energy assessment, the ES valuation would depend on the methods production, nutrient cycling, carbon sequestration, soil for- used. Therefore, it is important to know the strengths and mation, greenhouse gas emission, biological control of weaknesses of different methods used for a particular eco- pests and diseases, etc.; are the ES with tight coupling to system function assessment so that the researchers can the ecosystem processes and dynamics; and can be better make informed decisions. quantified by on-site field measurements whereas the cul- Hence, the aim of this review is to (a) provide an tural services can be better estimated by survey or ques- overview of key assessment methods in nutrient cycling tionnaire interview methods. ES valuation has been *Corresponding author. Email: email@example.com © 2014 Taylor & Francis 178 B.B. Ghaley et al. and carbon sequestration and (b) identify strengths and ‘carbon assessment methods*’ keywords to retrieve inves- weaknesses of the assessment methods and make sugges- tigations related to carbon sequestration and nutrient tions for improvement. cycling ES. Over 10,000 hits were available from the ISI Web of Science database and only studies that fulfilled the following criteria were included in this review: Methods (a) Diversity of studies on nutrient cycling and carbon In this study, literature published on the links between sequestration were maintained by selection of ecosystem processes and ES were compiled. Among the investigations in different terrestrial ecosystems ecosystem processes (Table 1), nutrient cycling and carbon (b) Methodologies applied were well described in sequestration process were the most common ecosystem details with a corresponding discussion of the processes investigated in diverse ecosystems like grass- methods land, freshwater, forest, shrubland, arable crops, aban- (c) Detailed descriptions were provided on trial loca- doned field and wetland. Given the high number and tion in terms of vegetation, soil type, location and diversity of the studies on nutrient cycling and carbon climatic information sequestration in different ecosystems, we chose to look (d) Results were provided in tabular formats instead at different methods used to study them. Literature search of graphs/bar charts to read the accuracy of the was carried out in ISI Web of Science with keywords like results under each investigation ‘ecosystem* service*’, ‘ecosystem* process*’, ‘ecosys- tem* function*’, ‘ecosystem* method*’, ‘ecosystem* good*’‘method* comparison*’ and the above keywords From the database, the most prominent methods used for were combined with ‘decomposition*’ and ‘mineralisa- determining nutrient cycling (decomposition and minera- tion*’, ‘soil organic carbon*’ and ‘soil carbon*’‘green- lisation) and carbon sequestration were compiled to pro- house gas*’ and ‘soil organic matter*’, ‘models*’ and vide an overview of methods and their descriptions. The Table 1. Ecosystem goods and services augmented by different ecosystem functions with examples. Ecosystem goods and services Ecosystem function Examples Provisioning 1 Food, feed, timber, Provision for plant growth/physical support Production of arable crops, pasture, timber and bio-energy supply bio-energy crops (Ghaley et al. 2014) 2 Raw material supply Provisioning of raw materials Timber for house construction (Ghaley et al. 2014) 3 Genetic resources Support diversity of plant genotypes Pool of resistance genes for control of pests and diseases (MEA 2005) Supporting 4 Primary production Support plant growth Support for growth of autotrophs 5 Soil formation Soil processes Weathering of rock fragments 6 Nutrient cycling Medium for storage, processing and transfer of Supply of NPK nutrients for plant growth nutrients (decomposition and mineralisation) (Sandhu et al. 2008) Regulating 7 Biological pest control Habitat for diversity of predators and pests Natural pest control dynamics in organic farming (Porter et al. 2009) 8 Water quality and water Filtration, buffering capacity and hydrological flow Drinking water quality for consumption and supply regulation (flood conservation of moisture (Ghaley et al. mitigation) 2014) 9 Erosion mitigation Formation of soil aggregates Reduction of erosion in steep slopes (Ghaley et al. 2014) 10 Global climate regulation Carbon sequestration, maintenance of atmospheric CO /O regulation and greenhouse gas 2 2 balance of gases and global temperature and regulation (CH and N O) (MEA 2005) 4 2 precipitation regulation 11 Global cycle of energy and Recycling of wastes and detoxification Human wastes are degraded/detoxified in soil matter flow and made available for recycling (TEEB 2010) Cultural 12 Recreation Platform for recreational activities Eco-tourism, mountain-climbing, hiking (MEA 2005) 13 Aesthetic value Social value preservation Landscape value, education and archaeological records (TEEB 2010) International Journal of Biodiversity Science, Ecosystem Services & Management 179 weakness and strengths of the methods were gleaned from Synthesis of methods to assess nutrient cycling different studies and reviews based on the discussions An earlier review on the methods of nutrient cycling provided. determination (Knacker et al. 2003) concluded that each method has its advantages and disadvantages and the use of one method over another will depend on the objective of the study. There is increasing emphasis on the need for Nutrient cycling consistency of methods with relevance for policymaking Nutrient cycling is a key ES that contributes to supporting in natural resource management. Since the ES provision is life on earth. Some of the prominent nutrient cycling spatially and temporally explicit, the field-level measure- processes include nitrogen fixation, phosphorus acquisi- ment methods should, as far as possible, reflect the differ- tion by mycorrhizal fungi and litter decomposition and ences to represent the ground reality. mineralisation. The increased availability of nitrogen Methods to assess nutrient cycling are compared through biological nitrogen fixation by rhizobium and based on ecological processes in operation, experiment phosphorus supply by arbuscular mycorrhizal fungi (dos cost and measurement end points (Table 3). With regard Santos et al. 2010; Gyuricza et al. 2010), are well docu- to ecological processes in operation, each method is mented nutrient cycling processes. Litter decomposition weighed against representativeness of the real field con- and mineralisation is an ecosystem process, resulting in ditions, integrating chemical, biological and physical breakdown of organic materials into their constituents by processes of organic matter breakdown. The decomposi- which nutrients are made available for nutrient cycling. tion rate is the result of the overall soil processes taking The key methods to determine litter decomposition and into account free access of soil fauna, terrestrial vegeta- mineralisation are as follows: tion quality and the environmental conditions prevalent at the trial site. Depending on the criteria chosen, some (a) Bait lamina method: The method consists of a methods have advantage over others and vice versa. For PVC strip (dimension 15–20 cm × 0.5 cm) with example, stable isotope and NIRS methods use the nat- 16 holes (dia. 1 mm), filled with bait material ural field conditions and reflect the actual decomposition (mixture of cellulose powder, bran flakes, agar- rates or mass loss at a particular location whereas litter- agar) and exposed to biogenic decomposition pro- bag and mini container method may alter the natural cess in the soil for measurement of the biological conditions due to microclimatic conditions induced by activity of the soil confining the litter material, affecting decomposition (b) Litter bag method: Assesses decomposition rates rate or mass loss. In contrast, litterbag and mini con- of organic materials based on plant litter loss tainer methods have an advantage in testing materials of enclosed in litter (mesh) bags and incubated in varied quality whereas there are limitations with stable situ or in meso or microcosms isotope and NIRS methods. For example, experiment (c) Mini container method: The method is a minia- set-up costs are minimum in stable isotope and NIRS turised version of litter bag technique (Eisenbeis methods whereas the cost is higher in litterbag and mini et al. 1999) and mini containers (MC) are poly- container methods. In contrast, stable isotope and NIRS ethylene boxes (volume ca. 1.5 m with diameter requires expensive equipment, whereas litterbag and mini container methods do not require such expensive 16 mm) containing organic substrates like litter, equipment. In case of measurement end points, bait straw and cellulose, closed on both ends by plastic lamina method is most disadvantaged among the meth- gauze of different mesh sizes (e.g 20 µm ods described because the bait lamina method does not −2.0 mm) (d) Near-infrared reflectance spectroscopy (NIRS) provide information on soil flora and fauna, mass loss, method: The information on types of bonds elemental analysis and decomposition rate. The reason is between the functional groups atoms in plant or that some investigations have reported that the number animal tissues can be obtained by spectroscopy, of baits consumed is indicative of soil macrofauna or based on principles of near-infrared region of elec- mesofauna activity and do not measure the activity of tromagnetic spectrum (750–2500 nm) the whole range of soil organisms contributing to (e) Stable isotope method: The application of stable decomposition rate. In terms of experience with different isotope techniques is particularly important in eco- methods, the ‘litterbag method’ is the most extensively system processes like decomposition of biogenic used. However, there are concerns that the decomposi- matter, mineralisation of nitrogen, functioning of tion effects in litterbag experiments are additive effects the mycorrhizal fungi, trophic relationships of two factors, viz., soil invertebrates present in the between soil animals, etc., which are difficult to litterbag and the litterbag mesh size. Often, the com- study in situ in the field bined effect is reported as soil invertebrate effect and hence an additional plot may be required to separate the effect of litterbag mesh size from the soil invertebrate Brief descriptions of the strengths and weaknesses of effect. different methods are summarised in Table 2. 180 B.B. Ghaley et al. Table 2. Summary of methods to assess nutrient cycling. Methods Strengths Weaknesses references 1 Bait lamina � Quick a nd inexpensive method � Results depend upon soil moisture André et al. (2009); Hamel et al. temperature, and number of strips (2007); Rożen et al. (2010) used 2 Litter bag method � Litter decomposition under different � The multiple litterbag size effect OECD (2006); Smith & Bradford combination of factors like litter confounds with the variable litter (2003); Villanyi et al. (2008) types (leaves, roots, stems, etc., of amount effect across studies Cornu et al. (1997); Crossley & vegetation), ecosystems (forest, making it difficult for result Hoglund (1962) agricultural land, grassland, etc.) and comparisons management practices (fertiliser, pesticide, tillage, crop rotation, etc.) � Simple, cheap and popular method and convenient to have required number of replicates 3 Mini container � Dual advantages of bait lamina and � Decomposition may increase Dunger et al. (2002); Keplin & method litter bag method in terms of higher because the material has to be Huttl (2001); Knacker et al. number of replicates cut into smaller size before (2003) exposure The maximum mesh size of 2 mm limits access to larger soil macro-fauna 4 Near-infrared � Non-destructive, rapid, cost-effective � Not popular in soil analysis due to Du & Zhou (2009); reflectance and reproducible method significant inorganic component Nduwamungu et al. (2009); spectroscopy in soil Reeves (2010); (NIRS) Wet samples obscure spectral information from the compounds of interest 5 Stable isotope � Easy to trace source and sink of The standard isotopic signature of Hyodo et al. (2010); Moore et al. methods different element cycles certain plant or animal tissue or (2004); Tiunov (2007) � Biological marker of matter and plant litter is difficult to determine energy flow and to evaluate the due to multiple sources of variation direction and rate of ecological processes Table 3. Comparison of assessment methods of litter decomposition and mineralisation with + and – indicating strengths and weaknesses of the method accordingly (adapted after Knacker et al. 2003). Criteria Bait lamina Litterbag Mini container NIRS Stable isotope 1. Relevance to ecological process Use in terrestrial ecosystem + + + + + Natural field conditions –– – ++ Relevance to local test material – ++ + + Tests of variable litter quality – ++ –– Access to soil fauna + + – ++ Data reproducibility – ++ + – 2. Experimental cost Equipment required + + + –– Experiment set-up cost + –– ++ Sample analysis cost + –– + – 3. Measurement end points Determination of decomposition rate – ++ + + Analysis of elemental composition – ++ + + Soil fauna – ++ ? + Soil microbial organism – ++ + + 4. Experience with technique – + –– – + – + – + – + – + – 5 /9 11 /3 9 /5 10 /3 9 /5 Soil carbon sequestration value for the welfare of human beings (Lal 2004). The major functions are as reservoir of major and minor plant Soil carbon is a natural capital, which provides multitude nutrients, soil aggregation for improved soil tilt and functions contributing to the economic and ecological International Journal of Biodiversity Science, Ecosystem Services & Management 181 reduced soil erosion, energy source for soil biota and viz., closed (steady state) or open (non-steady state) cham- buffers greenhouse gas emission from soil to the atmo- bers or by micrometeorological methods like eddy covar- sphere (Lal 2004). Soil carbon consists of two compo- iance technique consisting of ultrasonic anemometer and nents; soil inorganic carbon (SIC) and soil organic infrared gas analyser and the descriptions of the methods carbon (SOC) (Mishra et al. 2010). SOC is the major are provided below: constituent of soil organic matter (SOM) and forms the bulk of the soil component in different ecosystems across (a) Eddy covariance method is based on the principle biomes (Tornquist et al. 2009). SOC pool constitutes the that the atmosphere contains turbulent motions of largest component (1500 Gt.) in the biosphere; three times vertical movement of air, responsible for diffusion the terrestrial biomass pool and two times the atmospheric of atmospheric gases like CO . Such turbulent pool (IPCC 2007). SOC stock is affected, spatially and motions of air are sampled to assess the different temporally, in an ecosystem due to climate, terrestrial gases crossing a canopy-atmosphere interface by vegetation, soil mineralogy, soil management and the using statistical procedures for calculating vertical mass flux density. It is a measure of net CO interactions between these factors (Canadell et al. 2007). Past inventory of SOC dynamics has demonstrated that the exchange between the canopy-atmosphere by practice of clearing vegetation for agriculture and conven- micrometeorological methods with flux footprint tional production practices like intensive soil tillage have (the area of sampling) ranging from several hun- steadily increased the CO emission and other greenhouse dred metres to kilometres. The method is based on gases (CH and N O) concentration in the atmosphere, the physiology of the individual components in an 4 2 responsible for global warming effects (IPCC 2007). ecosystem and works best under non-undulating Carbon sequestration depends on the net balance of terrain, steady environmental conditions over large carbon fluxes and the assessment of the carbon budget of a sampling areas. Non-compliance to these assump- production system would require identification of carbon tions can amplify errors specially when deducing budget components (Smith et al. 2010). The gross uptake the flux measurements at different temporal scale of CO for photosynthesis in a field is given by the from days to years accumulation of gross primary production (GPP) of a (b) In closed chamber method, air sample is drawn production system (Luyssaert et al. 2010), a part of from a confined volume of air in the chamber for which is utilised for respiration by the plant itself for repair analysis by a connected infrared gas analyser, after and maintenance of existing plant tissues (autotrophic which the gas is returned to the chamber and net respiration) while the remaining portion is available for CO exchange is based on the rate of CO con- 2 2 synthesis of new plant organelles for growth and is called centration increase in the chamber placed over the net primary production (NPP). After accounting for loss of canopy or soil. In open chamber, the same princi- carbon in NPP, through decomposition process by soil ple is in operation except that the rate of CO fauna and microbes (heterotrophic respiration), the net concentration is based on the air flow gradient carbon available is called the net ecosystem productivity difference between the incoming air into the chamber and the air leaving the chamber after (NEP) (Ciais et al. 2010). The net ecosystem carbon budget (NECB) applies to net flux of carbon gain or loss equilibrium steady state is reached in the chamber in an ecosystem taking into account physical, biological headspace. Open and closed chamber methods are and anthropogenic sources and sinks and is given as more common in field experiments for net trace (Chapin et al. 2006; Smith et al. 2010). gas (CO ) flux measurements. A brief summary of strengths and weaknesses of the NECB ¼ NEE þ CO þ CH þ VOC þ DIC methods are provided in Table 4. þ DOC þ PC Synthesis of methods to assess carbon sequestration where NEE is the net ecosystem exchange of CO flux into the atmosphere, CO is net carbon monoxide flux, CH An earlier methodological review on carbon budget is the net methane flux, VOC is the net volatile organic assessment in ecosystem (Chapin et al. 2006) highlighted carbon flux, DIC is the net flux of dissolved inorganic the challenges posed by spatial and temporal differences carbon (leaching), DOC is the net flux of dissolved over short distances in terrestrial ecosystems. The review organic carbon (leaching) and PC is the net lateral transfer calls for the need for complementary approaches like of particulate carbon accounting for natural processes like literature review and use of models to triangulate the erosion, deposition by wind and water movement and field scale measurements to account for spatial and tem- anthropogenic activities like applying manure, crop har- poral difference in carbon fluxes across ecosystems. The vest and other processes like animal movement and loss research into various factors affecting greenhouse gas due to fire. The use of NECB over a large spatial scale is emission have pointed out that land use pattern, soil man- also called net biome productivity (NBP) (Chapin et al. agement and cropping systems have a significant role to 2006). NEE can be measured by chamber-based methods, play (Bernoux et al. 2006). The land use change is 182 B.B. Ghaley et al. Table 4. Summary of methods to measure carbon sequestration. Methods Strengths Weaknesses References 1. Eddy covariance � Robust in measuring fluxes at different � The flux measurement above the canopy Chapin et al. (2006); temporal scales from hours to years level may not be equivalent to the Luyssaert et al. (2009); Wu � Method is based on the physiology of actual course of gas flux across a soil- et al. (2010) the individual components in an plant system viz. during the night time ecosystem Build- up of CO due to respiration may � Any two ecosystems of interest can be not be recorded in the equipment compared for plant functional types placed at a certain reference height and composition, soil and crop from the canopy level, which can management regime underestimate the respiration during night time due to several reasons like insufficient turbulent mixing, false reading of the CO stored in the air space and the soil and outflow of CO from the canopy volume during night time 2. The chamber- � Low cost and flexibility to sample the � Needs a number of representative Savage et al. (2008); Tang based methods sites samples within the ecosystem to arrive et al. (2008); Zha et al. � Convenient to separate closed-spaced at a good estimate of flux rates (2007) treatments � The volume of air in the soil needs to be � Easy to differentiate fluxes from taken into account in closed chamber different ecosystem components systems to get a realistic flux estimate. However, soil air content is highly variable spatially and temporally and depends on soil type, vegetation; moisture status, etc., and such site- environment interactions affects flux estimation estimated to contribute 135 ± 55 Pg C due to decomposi- covariance technique is more capital intensive and is suita- tion of vegetation and mineralisation of humus or SOC ble for large non-undulating areas while chamber-based during the period 1850–1998 (Lal 2004). Adoption of methods are more suited for field experiments on smaller mitigation measures like no-till agriculture can help restore plots and is less expensive compared to eddy covariance. SOC pool by net gain of atmospheric CO into the soil and The long-term carbon stock change is better estimated by this process is called carbon sequestration (IPCC 2007). In use of process-based models, which can take into account reduced tillage cropping systems (e.g. conservation agri- various inputs of climate, management, farming system, culture, agroforestry), there are three main mechanisms in etc., over several decades at national and global scale. The operation, which help to conserve SOC. First, reduced key input into process-based models is the soil carbon data tillage reduces disruption of soil aggregates and avoids from a long-term experiment, which can be used to calibrate flush of microbial activity (Dameni et al. 2010), which and validate the model outputs to enhance robustness and protects SOM from decomposition. Second, reduced til- scientific credibility. The choice of a particular model will lage not only decreases soil erosion due to cultivation but depend on the objective of the study as all models are also avoids mixing of fresh residues into the soil, thus developed with a particular objective in mind. reducing decomposition processes. Soil cover is another significant factor that helps reduce soil erosion. At the global scale, conservation agriculture is estimated to Discussion −1 −1 store 0.1–1.3 t C ha year and can be practised in Soil-based ecosystem function assessment 60% of the arable land under agriculture (Lindstrom et al. 1998). The carbon sequestration potential in soils Nutrient cycling and carbon sequestration ES have signif- can differ and can be higher in degraded soils than in non- icant importance at the farm/field scale as well as at the degraded soils (Lal 2004). regional and global scale. At the farm/field scale, nutrient Given the importance of soil carbon in influencing the cycling and carbon sequestration are vital to maintain the on-site crop productivity and off-site GHG emission and productive capacity of the soil whereas at the regional or climate change, there is lot of emphasis on both soil carbon global scale, the two ES have equal importance in main- flux and stock determination with various methods. The taining the biogeochemical cycle and thereof mitigating short-term soil carbon measurements, viz., carbon fluxes adverse climatic effects. These two ES are dependent on can be better measured by dedicated field investigations the aboveground and belowground flora and fauna diver- with eddy covariance and chamber-based methods. Eddy sity. Reduction in above ground plant diversity can result International Journal of Biodiversity Science, Ecosystem Services & Management 183 in a variety of effects on belowground diversity and abun- conflicting study outcomes. Hence, determining humus dance of soil biota (Hooper et al. 2000), which in turn can stoichiometry could be a way forward to assess the gaps affect soil function or ES. Thus, improving the capacity of in building up SOM, with resultant effects on nutrient soils to provide adequate ES via soil biota functions can cycling and carbon sequestration. enhance biomass production. These ES provided by soil, Nutrient cycling influences nutrient dynamics in eco- support crop production by providing growth media for system, inter-specific interaction and ecosystem stability, seeds, aeration, plant support, nutrients (timely supply of thereby affecting ES provision (Ballantyne et al. 2008). nutrients and biological N inputs through fixation), water An ecological stoichiometry assessing different element and accumulation of carbon (De Groot et al. 2002; Sandhu and the ratios among them offers a potential approach to et al. 2012). Biological components of soils include assess effect of human activities on ES. The implications macro-, meso- and micro-fauna as well as micro-flora of stoichiometric ratios are quite explicit in agro- carry out ecosystem functions in the form of supplying ecosystems relating to crop quality and productivity, crop nutrients, biological control, organic matter turn over, pest populations, natural pollinators and control of pests maintaining soil structure and support the provision of and diseases and nutrient export. ES by soil (Gupta et al. 2010; Sandhu et al. 2010b). Although the application of N fertilisers may increase These ES are the result of complex interactions between the protein and N content in crop yield (Townsend et al. biotic (living) and abiotic (chemical and physical) compo- 2003), the high C: nutrients and N: nutrients ratio reduces nents of soil ecosystems through the universal driving the overall nutritional content of the crop, degrading the forces of matter and energy (De Groot et al. 2002). To ES value (Loladze 2002). For the crop pests and predators, capture the benefits of soil biological activity, a better nitrogen is the limiting nutritional factor and altered C:N understanding of the linkages between soil biota and eco- ratio can have both the positive and negative effects. High system function is required. plant C:N ratio can create a poor nutritional environment for the herbivores, which may reduce the negative effect of crop damage by pests but will also affect the growth and C:N stoichiometry ratios for ES assessment development of beneficial insects for control of pests (Wu Ecological stoichiometry has gained immense significance et al. 2010). In contrast, herbivores may increase their leaf after the work of Alfred Redfield (Cleveland & Liptzin feeding rates to meet the N deficit posed by high C:N ratio 2007), for developing Redfield Ratio, a correlation caused by increased CO levels. Such shifts in feeding between elemental composition of organisms and inor- habits causing increased crop damage without any adverse ganic nutrient in sea as 106C:16N:1P. The work of impact on their fitness would call for modified pest man- Sterner and Elser (Elser et al. 2010) further reinforced agement strategies under high C:N ratio context, demand- the prevalence of C:N:P ratios and stoichiometric home- ing revisiting of economic threshold levels for pesticide ostasis in plants and how it is being influenced by the local applications (Coviella et al. 2002; Reddy et al. 2004). environment. The relative composition of different organ- If the C:N ratio is decreased with fertiliser applica- isms differing in elemental stoichiometry has implications tions, there is tendency of pest population to increase on the food web interactions, contributing to the biogeo- (Cisneros & Godfrey 2001). However, there is no clear understanding on the effect of N deposition on the pest chemical cycles at global scale. Similar cascading effects population in the agricultural fields due to limited field have been widely studied in aquatic food web chain where studies. The biological control of pests is a central element changes in N:P ratio in algae effects the succeeding ani- of sustainable agriculture and altered stoichiometry in crop mals in the food chain (Vanni et al., 2011). The anthro- pogenic activities are related to disturbance of elemental plants may influence the predator performance. The high composition in nature, resulting in CO emission into C:N ratio causing reduced pest activity may improve the atmosphere, carbon sink in soil, nitrogen loading in arable control of pest populations by predators. Hence, the slower fields, etc., and insights into ecological stoichoimetry can growth of pests may allow for greater control by predators help unravel the role of elemental ratios in ecosystem as proposed by slow growth-high mortality hypothesis structure and processes for sustained ecosystem function (Benrey & Denno 1997). to generate ecosystem goods and services. To illustrate the The stoichiometric ratios can exert effects on pollina- significance of ecological stoichiometry framework on tors and pathogens in agriculture field affecting crop yield. nutrient cycling and carbon sequestration ES, the stoichio- High C:N ratios can render pollen as less preferred food metry of SOM can be used as an example. The importance for pollinators altering the pollinator foraging behaviour of SOM for nutrient cycling and retention and carbon and decreasing pollination efficiency and seed set and sequestration has culminated in numerous studies but thereof crop production (Poulton et al. 2001). there are conflicting reports from different production Susceptibility of plant disease due to change in stoichio- systems, management regimes and climatic conditions to metric ratios of host plant have been demonstrated in few build up SOM (Rumpel 2008). There is increasing under- studies (Hoffland et al. 2000; Solomon et al. 2003). For standing that the recalcitrant portion of SOM (humus) has example, high C:N ratio of litter slows down decomposi- a fixed C:N:P:S ratio (Kirkby et al. 2011) and the limited tion and crop residues are retained for longer period of supply of one of the nutrients is the potential reason for time, which might harbour disease inoculum and 184 B.B. Ghaley et al. 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International Journal of Biodiversity Science, Ecosystem Services & Management – Taylor & Francis
Published: Jul 3, 2014
Keywords: ecosystem functions; litter decomposition; mineralisation; assessment methodologies; stoichiometry
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