土食性大型土壤动物转化土壤有机碳的14C示踪法应用研究进展

土壤有机C是维持全球C平衡过程中的重要C库,其降解和转化在地球化学循环中占有重要地位。大型土壤动物对土壤有机C的稳定性起着重要的调控作用。14C示踪技术由于在测定目标化合物的转化速率和定位代谢产物和残留物分布上的优势,近年来在土壤有机C稳定性研究中得到了广泛应用。本文综述了3种典型土食性大型土壤动物,白蚁(Isoptera:Termitidae)、甲虫幼虫(Coleoptera:Scarabaeidae)和蚯蚓(Oligochaeta:Lumbricidae),对土壤稳态有机C降解转化的14C示踪研究结果及相关的微生物作用。食土白蚁和甲虫幼虫的极端碱性(pH10～12.5)肠道段和肠道内的特殊蛋白酶的共同作用促使这两种动物可以选择性地降解和矿化腐殖酸中的稳态多肽等组分,进一步提高腐殖酸的腐殖化程度。食土蚯蚓体内含有高活性的纤维素酶,能促进纤维素的降解。虽然木质素在蚓粪中能发生降解,蚯蚓也能降解植物树叶,但是食土蚯蚓能否降解或选择性降解稳态土壤有机C的研究还极少。大型土壤动物肠道内含有大量微生物及酶,这些微生物在土壤动物降解和腐殖化有机C的过程中所起的具体作用如何以及这些酶的来源还不是很清楚。文中总结了目前对大型土壤动物转化土壤有机C认识上的不足,并对一些优先研究方向提出了建议。     

Transformation and mineralization of 14C-labeled cellulose, peptidoglycan, and protein by the soil-feeding termite Cubitermes orthognathus

 We performed feeding trials with the soil-feeding termite Cubitermes orthognathus using soil spiked by uniformly ¹⁴C-labeled preparations of cellulose, peptidoglycan, protein, and bacterial cells (Bacillus megaterium and Escherichia coli). When incubated in soil for 8 days in the absence of termites, cellulose and peptidoglycan showed low mineralization rates (0.5% and 0.2%, respectively). However, when termites were present, their mineralization rates strongly increased (21.6% and 30.6%, respectively). The mineralization rate of protein was 12.4% in the control soils and increased to 36.2% in the presence of termites. Mineralization of bacterial cells in control soils occurred in two phases (rapid mineralization during the first 4–5 days and stabilization thereafter). When termites were present, the rates of mineralization of bacterial cells increased and the stabilization phase was abolished. In all cases, radiolabel accumulated in the termites and the solubility of the labeled compounds located in the gut increased strongly. Mineralization was accompanied by transformation of residual carbon from the humic acid fraction to the fulvic acid fraction during gut passage. High-performance gel permeation chromatography demonstrated a strong shift in the size distribution of the residual carbon from high-molecular-weight towards low-molecular-weight molecules in the gut of termites and an accumulation of small molecules in the termite bodies. The present study provides strong evidence that structural polysaccharides of plants and bacteria and microbial biomass are carbon and energy sources for soil-feeding termites. Received: 29 May 2000     

Subterranean termites: regulators of soil organic matter in the Chihuahuan Desert

Soil organic matter and the abundance of subterranean termites were measured at 89 locations spaced at 30-m intervals from the bottom to the top of a small desert watershed. There was no correlation between soil organic matter content and topographic position on the watershed. Analysis by autocorrelogram demonstrated that the soil organic matter content was randomly distributed on the watershed. There was a highly significant negative correlation between termite abundance and soil organic matter, r=–0.97. Soils characterized by horizon in soil pits within each vegetation type (soil type) showed some relationships to erosion-deposition areas on the watershed, with surface organic matter contents varying between 3.4% in the playa basin where termites were absent to 0.4% in a sparse shrubland on erosional soils. In the northern Chihuahuan Desert, subterranean termites appear to be responsible for most of the variation in soil organic matter.     

Theoretical approaches to modelling the dynamics of soil organic matter

Two theoretical concepts of the formation of the soil organic matter (OM) system are considered as a methodological basis for the mathematical simulation of its dynamics. In the theory of physical protection of OM without the formation of humic substances with the physical fractionation methodolody, the main focus is on the functional parameters of the OM components separated by their mineralization rates. Here, two conceptual disadvantages are noted: (a) neglect of the specificity of OM transformation in organic soil horizons, where humified OM resistant to mineralization is formed, which cannot be explained by this theory; and (b) consideration of the soil microorganisms as a unified undifferentiated complex. In classical humification theory, a number of humification stages are considered with the respective communities of decomposer organisms that mineralize OM and transform biota products into humic substances. The silvicultural concept of humus forms was found to be effective and suitable for a wide range of natural conditions with the use of this theory for OM simulation. There is a general shortcoming to both approaches: protection theory has no parameters of recalcitrant OM formation from other fractions; in humification theory, the quantitative humification parameters under the effect of soil fauna have yet to be sufficiently substantiated. The values of the turnover time for active, intermediate, and recalcitrant OM are revised. The importance of theoretically substantiating the structural and functional organization of OM for its dynamic modelling is emphasized.     

Influence of blood meal organic fertilizer on soil organic matter: A laboratory study

Blood meal is an organic nitrogen (N) fertilizer containing about 10–13% organic N. The fertilizer is mainly composed of hemoglobin which is characterized by the presence of a prosthetic group containing iron (Fe). Knowledge about the influence of blood meal on soil organic matter and soil fertility is now limited. In this work, blood meal was incubated in the soil for one year and the evolution of the organic matter was followed at time intervals by use of isoelectric focusing (IEF) and humification parameters. The mineralization of the total N and the availability of some mineral nutrients (Fe in particular) were also followed. The results show that only a part (about 75%) of the organic carbon (C) and of the organic N mineralized and that the remaining C was transformed into humified compounds. The availability of the Fe increased during the incubation, probably due to the progressive degradation of the prosthetic group and the successive chelation of the Fe from the humic substances.     

Qualitative and quantitative soil organic matter estimation for sustainable soil management

Purpose

The aims of this paper were to review tools and methods for qualitative and quantitative evaluation of soil organic matter (SOM) coming from diverse egzogenic sources for effective soil management, and to introduce a new approach to predict dynamics of SOM transformations, especially humification, as a key process in the formation of humic substances (HSs).

Materials and methods

A review of existing literature is presented on tools and methods for qualitative and quantitative assessment of organic matter in soil originating from various sources for reasonable soil management, attempting to provide a better understanding of the advances in organic matter transformations and new research directions for modeling. Diverse tools and methods for qualitative and quantitative evaluation of organic matter in soil coming from diverse sources have been adopted so far to express transformation processes.

Results and discussion

For the qualitative analysis of SOM and humic acids (HAs), the analytical techniques are applied, e.g., HPSEC, NMR, and ESI-FTICRMS. The quantitative analysis is done through the following parameters: humification index (HI), humification degree (HD), and humification rate (HR). These analyses indicated that because of lack of reliable data from sufficiently long-term experiments, mathematical modeling may be applied as a numerical tool for quantitative estimation and prediction of humification of SOM. The effective soil management should include soil properties as well as different functions: food production, nutrient and water cycling, storage, filtrating, buffering, biological habitat, gene pool, source of raw materials, climate regulations, heritage, platform for man-made structure. The soil utility value should be evaluated through the SOM qualitative and quantitative analysis of organic carbon and total nitrogen. Knowledge about dynamics of SOM transformations is essential, particularly in the context of stability and efficiency of different sources of organic matter applied into soil. A qualitative understanding of SOM dynamics transformations along with modeling for quantitative assessment of HS formation should be used to develop sustainable soil management. The modeling may be considered as a tool for predicting SOM humification dynamics and consequently the formation of HSs from the diverse sources. The existing archival data from a long-term experiment may be used to build and calibrate the reliable mathematical model of SOM humification.

Conclusions

Managing of SOM remains a sound basis for maintaining soil in a good condition for optimizing productivity. The development of land management strategies to optimize both the increase of soil organic carbon levels and the recycling of nutrients from SOM needs to be a priority. This should include policy makers and other users as well.

     

Degradation and humification of maize straw in soil microcosms inoculated with simple and complex microbial communities

Microbial communities are responsible for soil organic matter cycling and thus for maintaining soil fertility. A typical Orthic Luvisol was freed from organic carbon by thermal destruction at 600°C. Then the degradation and humification of ¹⁴C‐labelled maize straw by defined microbial communities was analysed. To study the role of microbial diversity on the humification of plant material, microcosms containing sterilized soil were inoculated with a natural microbial community or with microbial consortia consisting of bacterial and fungal soil isolates. Within 6 weeks, 41 ± 4% of applied ¹⁴C‐labelled maize straw was mineralized in the soil microcosms containing complex communities derived from a soil suspension, whilst the most efficient communities composed of soil isolates mineralized less than 35%. The humification products were analysed by solution state ¹³C‐NMR‐spectroscopy and gel permeation chromatography (GPC). The analyses of humic acids extracts by solution state ¹³C‐NMR‐spectroscopy revealed no difference in the development of typical chemical functional groups for humic substances during incubation. However, the increase in specific molecular size fractions of the extracted humic acids occurred only after inoculation with complex communities, but not with defined isolates. While it seems to be true that redundancy in soil microbial communities contributes to the resilience of soils, specific soil functions may no longer be performed if a microbial community is harshly affected in its diversity or growth conditions.     

Transformation of the organic matter in an agrogray soil and an agrochernozem in the course of the humification of corn biomass

The transformation of the organic matter in the course of corn residues humification in an agrogray soil and an agrochernozem was studied in long-term experiments using the method of solid-phase ¹³C-nuclear magnetic resonance spectroscopy. The humification of the plant residues was found to be accompanied by a decrease in the content of the O-alkyl fragments comprising polysaccharides and polypeptides, an increase in the unsubstituted alkyds content, and by the relative accumulation of aromatic fragments and carboxyl groups. The most strongly transformed pool of the organic matter, as compared to the initial plant residues, was the humic acids with their maximal content of aromatic and carboxyl functional groups and the minimal content of O-alkyls. The chemically stable aromatic fragments were concentrated not only in the pool of humic acids; their content was 64–89% of the pool of the aromatic fragments identified in the soil organic matter. Therefore, to assess the stable pool, the distribution of the functional groups is necessary to be analyzed not only in the humic acids but also in the whole soil organic matter.     

The response of soil organic matter to manure amendments in a long-term experiment at Ultuna, Sweden

In a long-term field experiment started in 1956 on a clay loam soil at Uppsala, Sweden, changes of organic carbon in the topsoils receiving various organic amendments at the rate of 200 kg C ha^'1 year^'1 were studied to determine soil organic matter characteristics, variations of δ¹³C in the soil and to estimate a carbon balance. Fallow and mineral fertilizer without N led to a significant decrease of soil organic matter (SOM) in the soil, green manure maintained the SOM content, and animal manure and peat increased the SOM content significantly. The stable portion of the added organic materials after 37 years of continuous input was 12·8, 27·3, and 56·7%, for green manure, animal manure and peat, respectively. This was reflected by half-lives of organic carbon originating from the amendments between 3·0 (green manure) and 14·6 years (peat). The isotopic composition of SOM changed both due to mineralization (continuous fallow) and the addition of amendments is topically different from soil humus (green manure, animal manure). The isotopic effect was used to calculate the percentage of carbon derived from animal manure present for the year 1993. This value (55·4%) was larger than that derived from the carbon balance, which indicated a priming effect of the animal manure on the initial soil humus. Mineralization of microbially available organic substances led to an increase in the degree of humification on plots not receiving organic amendments. Adding peat and animal manure resulted in a decrease of the humification index due to the continuous input of poorly humified material. The extinction ratio (E₄/E₆) and ratio of fulvic acid to humic acid changed considerably in the peat treated plots. Fourier transform infrared (FTIR)-measurements of the extracts showed that peat characteristics can be detected in peat treated soils. The other amendments did not alter the characteristics of the extractable humic substances.     

Modelling long‐term carbon dynamics in soils reconstituted with large quantities of organic matter

In urban conditions, the widescale availability of organic matter to be recycled and the necessity for soil reconstitution (Technosol) has led to the input of very large quantities of organic matter (up to 50% v/v). The long‐term degradation of these large quantities of organic matter in the soil is not well known. We monitored, over a 60‐month period, the total carbon (C) content and the particulate and biochemical fractions of reconstituted soils placed in 600‐litre boxes under natural conditions. The top layer was a sandy loam amended with 20 or 40% of sphagnum peat or organic compost (sewage sludge, wood chip compost or green waste compost) lying on a layer of sandy loam. We measured C mineralization over time under controlled conditions and built a long‐term model to simulate carbon dynamics where exogenous organic carbon (EOC) was divided into two biodegradable compartments. The model yielded the proportions of EOC that either resisted degradation or contributed to soil organic C storage by mineralization and/or humification. Organic matter degradation was linked to its maturity and to its contents in certain particulate and biochemical fractions but was independent of how much of a given organic matter was introduced. We found a good correlation between the degradable organic compartment and the lignin and cutin‐like, hemicellulose and cellulose‐like fractions larger than 1 mm. The model showed that a large part of initial EOC was still present in the soil after 5 years in a potentially biodegradable but resistant compartment. The degradation of that compartment by mineralization or humification is therefore expected to take longer.     

Changes in soil properties and humic substances after long‐term amendments with manure and crop residues in dryland farming systems

After 16 years of periodical applications of either farmyard manure or crop wastes at two levels of mineral N fertilization to a Calcic Haploxeralf in the semiarid central Spain, we found significant changes in chemical fertility levels and in the concentration, chemical composition, and carbon mineralization rates of soil organic matter (SOM). The changes in SOM quality were related to significant improvements of soil physical properties, mainly aggregate stability and water retention. Such changes were related to the increased concentration of humic colloids in soil, the mineral N dose, and the type of organic matter applied. When compared with the control plots, the organic matter accumulated in the amended plots tended to be less transformed, and its total concentration and humification degree decreased with increasing external N‐inputs. Humic acids from the amended plots showed a more marked aliphatic character (mainly after N addition) than those from control plots. Farmyard manure led to a significant improvement of soil physical properties, but had a comparatively small effect in promoting biodegradation and humification of crop wastes. This could be due to the high biological stability of the manure used which, in semiarid Mediterranean fields, usually leads to an accumulation of little transformed SOM.     

Termites promote soil carbon and nitrogen depletion: Results from an in situ macrofauna exclusion experiment,Peru

We present results from one of the first in situ soil termite exclusion experiments using translocated soil that was not colonised by termites prior to the experiment. Macrofauna were excluded or included using fine (0.3 mm) and coarse (5 mm) mesh, respectively. We found that termites were the most dominant macrofauna in the macrofauna-included samples throughout the sampling period. Additionally, C and N depletion rates were consistently higher in samples with macrofauna than without macrofauna despite the seasonal decline of termites at the start of the wet season. This suggests that the presence of termites in soil promotes C and N depletion that may be linked to the passage of soil through the termite gut and the affect termites have on bioturbation and nutrient distribution.     

Effect of cultivation on chemical characteristics and respiratory activity of crusting soil from Mazowe (Zimbabwe)

Abstract

Clearing and cultivation in crusting soils from Mazowe (Zimbabwe) has lead to severe changes in most physico‐chemical characteristics related to the concentration and distribution patterns of plant nutrients and to the total amount of soil organic matter. Nevertheless, the concentration of the different humus fractions showed lower intensity changes, as did the mineralization rates of the organic matter. The most significant effects of cultivation on the soil chemical characteristics coincided with those considered to favor clay dispersion and crusting phenomena, including generalized desaturation of the exchange complex and losses of divalent ions with a potential bridging effect between soil particles. Concerning the soil organic matter, the humic acid tended to concentrate in the cultivated soils as a probable consequence of selective biodegradation of the other humic fractions. The composition and activity of soil humus suggest low‐performance organo‐mineral interactions: in these soils the active turnover of the plant wastes is not regulated by intense physico‐chemical interactions with the soil mineral fractions, or by physical encapsulation of organic particles. In consequence, the mineralization rates were relatively constant in the soils studied and unrelated to soil organic matter concentration. The results suggests that there is a possibility to revert the early degradation stages of these soils through a rational management of suitable amounts of crop wastes.     

Consistent effects of eastern subterranean termites (Reticulitermes flavipes) on properties of a temperate forest soil

Termites inhabit a large portion of land covered by temperate forests. Climate warming and urbanization will likely extend their range and increase their densities in these ecosystems but, compared to their tropical counterparts, little is known about their effects on soil properties and processes. If temperate termites have the strong ecosystem engineering effects of tropical termites, then knowledge of their ecology and impacts will be vital for predicting how temperate systems respond to environmental change. We investigated how feeding and tunneling by the eastern subterranean termite, Reticulitermes flavipes, affected wood decomposition and soil properties under decaying wood. Twelve laboratory microcosms filled with mineral soil and with wood blocks of four common temperate tree species, received R. flavipes soldiers and workers at field densities, with an additional five termite-free microcosms serving as controls. After 25 weeks, the effects of termites on wood mass loss, and on carbon and nitrogen dynamics, differed across tree species, yet their effects on soil properties were consistent regardless of wood type. Microbially-available carbon in soil was 20% higher with termites and soil moisture content 20% lower. Soil pH was more acid with termites and their effects on soil microbial biomass were positive but non-significant. These soil responses were consistent regardless of the wood species, suggesting that termite effects on soil are dictated largely by their activity within the soil matrix and not by their feeding rate on specific wood substrates. These results are among the first to quantify the effects of temperate forest termite activity on soil properties, demonstrating the potential for these termites to shape biogeochemical cycling and plant communities through their alteration of the soil environment.     

Humification-mineralization pyrolytic indices and carbon fractions of soil under organic and conventional management in central Italy

The aim of this study was to evaluate chemical changes in soil organic matter (SOM) in organically and conventionally managed fields, using pyrolytic indices and the extraction of different carbon fractions. Pyrolysis-gas chromatography (Py-GC) was used to study structural changes in SOM, whereas the different soil extractions gave a fractionation of C forms. Organic management increased both humic and labile C forms (microbial biomass C and water soluble organic C). A significant positive relationship was found between the living SOM fraction, expressed as microbial biomass/total organic C ratio (MBC/TOC) and humification rate. A negative relationship was found between the pyrrole to phenol ratio (O/Y) and total extractable C (TEC).An opposite trend has been observed for the second pyrolytic index (N/O), which represents the mineralization of fresh organic matter. Mineralization was higher in organically managed soil, probably because of consistent input of fresh material to the organic field. Carbon fraction pools and pyrolytic indices provided complementary indications of SOM quality under organic and conventional management.     

Evolution of Humic Fractions in Calcimagnesic Soils And Brown Isohumic Soils Amended With Compost and Manure

The composting process of organic wastes consists of the bioconversion of biologically unstable wastes to stabilized products for application to soils. This study was carried out on different soils (brown isohumic soils in Sfax region in southern Tunisia and calcimagnesic soils in Korba region in northern Tunisia). The mineralization process of organic matter is more important in isohumic soils, showing less tenors of fine fractions than in calcimagnesic soils. The study of different humic fractions in amended soils shows that humic acid tenors and the polymerization rates are linked to soil nature. Calcimagnesic soils ensure a more important humification than isohumic soils.     

Effect of different management strategies on soil quality of citrus orchards in Southern Italy

Twenty‐six soil samples were collected from 13 paired orchards (organically vs. conventionally managed) homogeneous for age, rootstock and cultivars, belonging to the Eastern Sicily Organic Citrus farm Network. The soil quality was evaluated by chemical and biochemical indicators. The total organic C, humification parameters and isoelectric focusing of extracted organic matter were measured to quantify the size of relevant soil C pools. In addition, C turnover was evaluated by determining microbial C mineralization, C microbial biomass and by calculating the mineralization and metabolic quotient (qCO₂). The results obtained demonstrated that organic citrus soils were characterized by a general increase in all the organic matter pools, which means a greater C supply for soil metabolic processes. This observed trend did not directly influence the organic matter turnover, indicating that the organic approach could act as a soil C‐sink. The soil microflora of organically managed soils showed an improved efficiency in use of energy and organic resources, corresponding to an increased ability of soils under organic management to sustain biological productivity in the long term.     

Composting And Humification

The development of humic substances in the course of four composting processes was monitored quantitatively, recording both relative and absolute contents. Relative data showed contrasting results if the humic substances (HS) were related to the dry matter (d.m.) or to volatile solids (VS). Humic substances were apparently formed because of a concentration effect due to organic matter degradation. If absolute contents were considered, a decrease in the humic substances was observed, above all in the early stages of the process, due, probably, to degradation of the organic material, such as proteins, carbohydrates and lipids, coextracted with the humic substances. Processing of the data in respect of humic substance content over 13 composting processes and one study on the degradation of plant residues in soils, confirmed that no net humic substances are formed during composting and that the humification should be interpreted merely as degradation of the organic matter associated with the humic substances, after uncovering what is known as the core of the humus. The concept of humification during the composting process therefore needs to be reviewed, bearing in mind that neither the method commonly used for humic substance extraction nor the relative results obtained enable it to be interpreted satisfactorily.     

Chemical studies on soil humic acids

Seventeen samples of soil humic acids, two fractions of soil fulvic acid sample, and several related compounds such as lignin, tannin, flavonoid and artificial humic substances were decomposed in conc. KOH solution at 180°C. Succinic acid, glutaric acid, phloroglucin, p-hydroxybenzoic acid, vanillic acid, protocatechuic acid, 3,4-dihydroxy-5-methoxybenzoic acid, and gallic acid were detected in the degradation products of humic acids. The amounts of these degradation products were discussed in relation to the degree of humification or the sources of the humic acid samples. Succinic acid also resulted from glucose, polymaleic acid, and the humic acid and humin prepared from glucose, but glutaric acid resulted only from glucose humic acid and glucose humin but not from glucose and polymaleic acid. Succinic acid and glutaric acid were supposed to result from the same structural portions in humic acids because of the very significant positive linear correlation between their amounts. p-Hydroxybenzoic, vanillic, protocatechuic, and 3,4-dihydroxy-S-methoxybenzoic acids were presumed to result mainly from lignin structure in humic acids. Soil humic acids yielded small amounts of gallic acid although the yields by hydrolysable tannins were in large amounts. The yields of above-mentioned degradation products from humic acids decreased with increasing degree of humification. Phloroglucin resulting from ftavonoids including condensed tannins were also found in the degradation products of humic substances. Its yield showed no linear correlation with RF value of humic acid, and is presumed to be rather related to the vegetation at the sites of soil sampling.     

Selective digestion of the peptide and polysaccharide components of synthetic humic acids by the humivorous larva of Pachnoda ephippiata (Coleoptera: Scarabaeidae)

In order to identify the potential nutrient and energy sources of humivorous beetle larvae, we carried out feeding trials with soil supplemented with specifically ¹⁴C-labeled model humic acids synthesized by peroxidase-initiated radical polymerization, using the cetoniid beetle Pachnoda ephippiata (Coleoptera: Scarabaeidae) as a model organism. Ingestion of soil by the larvae significantly increased the mineralization of humic acids labeled in their peptide (HA-*peptide) or polysaccharide components (HA-*peptidoglycan and HA-*chitin), whereas the mineralization of humic acids labeled in the aromatic components (HA-*catechol) did not increase significantly. Mineralization was accompanied by a reduction of residual radiolabel in the acid-soluble fraction and an increase in the humic acid and humin fractions of the fecal pellets. During the gut passage, the residual label in peptide or polysaccharide components was transformed into acid-soluble products, especially in the alkaline midgut. High-performance gel-permeation chromatography demonstrated that the changes in solubility were accompanied by large changes in the molecular weight of the residual material. The amount of radiolabel derived from the peptide and polysaccharide components recovered from the larval body and hemolymph was significantly higher than that derived from the aromatic component, which supports the hypothesis that humivorous beetle larvae selectively digest the peptide and polysaccharide components of humic substances, whereas the aromatic components of humic substances are not an important source of nutrients and energy. This is also the first experimental evidence that also chitin and peptidoglycan, the major structural polymers in fungal and bacterial biomass, can be protected from microbial degradation in soil by a copolymerization with phenols and might contribute substantially to the refractory nitrogen pool in soil organic matter.