Decomposition of litter produced under elevated CO2: Dependence on plant species and nutrient supply

We investigated the effect of CO₂ concentration and soilnutrient availability during growth on the subsequent decomposition andnitrogen (N) release from litter of four annual grasses that differ inresource requirements and native habitat. Vulpia microstachys isa native grass found on California serpentine soils, whereas Avenafatua, Bromus hordaceus, and Lolium multiflorum areintroduced grasses restricted to more fertile sandstone soils (Hobbs & Mooney 1991). Growth in elevated CO₂ altered litter C:N ratio,decomposition, and N release, but the direction and magnitude of thechanges differed among plant species and nutrient treatments. ElevatedCO₂ had relatively modest effects on C:N ratio of litter,increasing this ratio in Lolium roots (and shoots at high nutrients),but decreasing C:N ratio in Avena shoots. Growth of plants underelevated CO₂ decreased the decomposition rate of Vulpialitter, but increased decomposition of Avena litter from the high-nutrient treatment. The impact of elevated CO₂ on N loss fromlitter also differed among species, with Vulpia litter from high-CO₂ plants releasing N more slowly than ambient-CO₂litter, whereas growth under elevated CO₂ caused increased Nloss from Avena litter. CO₂ effects on N release in Lolium and Bromus depended on the nutrient regime in whichplants were grown. There was no overall relationship between litter C:Nratio and decomposition rate or N release across species and treatments.Based on our study and the literature, we conclude that the effects ofelevated CO₂ on decomposition and N release from litter arehighly species-specific. These results do not support the hypothesis thatCO₂ effects on litter quality consistently lead to decreasednutrient availability in nutrient-limited ecosystems exposed to elevatedCO₂.     

Climate and litter quality differently modulate the effects of soil fauna on litter decomposition across biomes

Climate and litter quality have been identified as major drivers of litter decomposition at large spatial scales. However, the role played by soil fauna remains largely unknown, despite its importance for litter fragmentation and microbial activity. We synthesised litterbag studies to quantify the effect sizes of soil fauna on litter decomposition rates at the global and biome scales, and to assess how climate, litter quality and soil fauna interact to determine such rates. Soil fauna consistently enhanced litter decomposition at both global and biome scales (average increment ~ 27%). However, climate and litter quality differently modulated the effects of soil fauna on decomposition rates between biomes, from climate‐driven biomes to those where climate effects were mediated by changes in litter quality. Our results advocate for the inclusion of biome‐specific soil fauna effects on litter decomposition as a mean to reduce the unexplained variation in large‐scale decomposition models.     

土壤动物对川西亚高山和高山森林凋落叶第一年不同分解时期N和P元素动态的影响

土壤动物对高寒森林凋落物养分元素动态具有重要影响, 但这种影响受控于凋落物质量及环境条件。为了解土壤动物对高寒森林凋落物不同分解时期凋落物中N和P元素动态的影响, 采用凋落物分解袋的方法, 于凋落物第一年分解的不同时期, 即冻结前期、冻结期、融化期、生长季节初期、生长季节中期和生长季节末期, 研究了3.00和0.04 mm孔径凋落物袋中川西亚高山和高山森林的代表性植物——康定柳(Salix paraplesia)、方枝柏(Sabina saltuaria)、红桦(Betula albosinensis)和岷江冷杉(Abies fargesii var. faxoniana)凋落物中的N和P元素动态特征。结果表明: 康定柳和红桦凋落物中的N元素呈现出释放—富集—释放的模式, 方枝柏、岷江冷杉凋落物中的N元素则表现为释放—富集模式; 凋落物P元素总体表现为释放模式, 但4种植物凋落物均在生长季节中期具有明显的富集过程; 从凋落物分解的第一年来看, 土壤动物明显促进了4种植物凋落物N的释放, 而抑制了P的释放; 不同时期土壤动物对凋落物中N和P释放量的影响存在显著差异, 且分别与正积温呈极显著正相关和极显著负相关关系; 相对于阔叶植物凋落物, 土壤动物对针叶植物凋落物中N和P元素动态的影响更为显著。这些结果为深入了解高寒森林生态系统土壤动物与凋落物分解等物质循环过程的相互联系具有重要意义。     

Effects of grassland-use on soil respiration and litter decomposition

 草地利用方式影响植被群落结构和土壤微环境, 制约草地生态系统碳循环。该文通过测定温带草原在放牧、割草、围封3种利用方式下湿润年(2012年)和干旱年(2011年)的凋落物产量、质量及其分解速率和土壤碳通量, 分析了草地利用方式对土壤呼吸和凋落物的影响, 探讨了凋落物对土壤呼吸的贡献机制。结果表明: 在干旱年份, 放牧样地土壤呼吸最大, 分别达到割草和围封样地的1.5倍和1.29倍; 在湿润年份, 割草样地土壤呼吸最大, 为309 g C·m^–2·a^–1, 明显高于放牧样地和围封样地。不论干旱年还是湿润年, 围封样地凋落物产量都大于放牧样地和割草样地。3种利用方式下湿润年土壤呼吸和凋落物分解均比干旱年增强。因此, 水分是温带草原植物生长和生态系统碳循环的主要限制因子, 草地利用方式则显著影响凋落物生产和分解。进一步分析表明, 经过两年的分解, 同一样地内凋落物质量C:N下降, N含量和木质素:N升高, 土壤呼吸与凋落物产量、凋落物分解速率以及木质素:N正相关, 而与凋落物C:N负相关。     

草地利用方式对土壤呼吸和凋落物分解的影响

草地利用方式影响植被群落结构和土壤微环境, 制约草地生态系统碳循环。该文通过测定温带草原在放牧、割草、围封3种利用方式下湿润年(2012年)和干旱年(2011年)的凋落物产量、质量及其分解速率和土壤碳通量, 分析了草地利用方式对土壤呼吸和凋落物的影响, 探讨了凋落物对土壤呼吸的贡献机制。结果表明: 在干旱年份, 放牧样地土壤呼吸最大, 分别达到割草和围封样地的1.5倍和1.29倍; 在湿润年份, 割草样地土壤呼吸最大, 为309 g C∙m^-2∙a^-1, 明显高于放牧样地和围封样地。不论干旱年还是湿润年, 围封样地凋落物产量都大于放牧样地和割草样地。3种利用方式下湿润年土壤呼吸和凋落物分解均比干旱年增强。因此, 水分是温带草原植物生长和生态系统碳循环的主要限制因子, 草地利用方式则显著影响凋落物生产和分解。进一步分析表明, 经过两年的分解, 同一样地内凋落物质量C:N下降, N含量和木质素:N升高, 土壤呼吸与凋落物产量、凋落物分解速率以及木质素:N正相关, 而与凋落物C:N负相关。     

Highly consistent effects of plant litter identity and functional traits on decomposition across a latitudinal gradient

Plant litter decomposition is a key process in terrestrial carbon cycling, yet the relative importance of various control factors remains ambiguous at a global scale. A full reciprocal litter transplant study with 16 litter species that varied widely in traits and originated from four forest sites covering a large latitudinal gradient (subarctic to tropics) showed a consistent interspecific ranking of decomposition rates. At a global scale, variation in decomposition was driven by a small subset of litter traits (water saturation capacity and concentrations of magnesium and condensed tannins). These consistent findings, that were largely independent of the varying local decomposer communities, suggest that decomposer communities show little specialisation and high metabolic flexibility in processing plant litter, irrespective of litter origin. Our results provide strong support for using trait-based approaches in modelling the global decomposition component of biosphere-atmosphere carbon fluxes.     

Interactions between atmospheric CO2 enrichment and soil fauna

We have reviewed the responses of soil fauna to increased concentrations of atmospheric CO₂ and the consequent climate change. These will affect several attributes of animal populations and communities including their density, biomass, diversity, activity, rates of consumption, life history parameters and migration ability. Changes in the quality and quantity of litter and global warming are the main factors which are expected to modify soil fauna. Although changes have been observed in several attributes of the soil fauna as a consequence of increased concentrations of atmospheric CO₂, no general trend which might allow to the prediction of a general pattern of response has been identified. Because of the complexity of the biological mechanisms and the synergetic action of several factors, the few resulting responses reported in the literature are inconclusive. However, some aspects of the situation deserve more attention. These include the consequences of (1) changes in the food resources for soil fauna in the litter layer and in the rhizosphere, (2) the consumption of low quality litter by the macrofauna, (3) the change in life span in response to temperature elevation, (4) the enhancement of earthworm burrowing activity and (5) the changes in community composition arising because of specific differential resistance to adverse conditions. This revised version was published online in June 2006 with corrections to the Cover Date.     

土壤动物与N素循环及对N沉降的响应

以主要的生态过程之一——N循环为对象,论述了土壤动物不仅对凋落物的分解有重要影响,而且在N素矿化和植物对N的吸收过程中也起着重要作用。同时,日益严重的全球变化问题之一——N沉降对土壤动物的多样性及其在生态系统中的功能构成了极大的威胁。另还对土壤动物与N循环研究的方法、土壤动物在N循环过程中的作用机制、热带地区的需求及N沉降下土壤动物的响应作了探讨,并提出,开展大尺度的专类研究及长期定位研究成为下一步研究的需要。     

C,N and P fertilization in an Amazonian rainforest supports stoichiometric dissimilarity as a driver of litter diversity effects on decomposition

Plant leaf litter generally decomposes faster as a group of different species than when individual species decompose alone, but underlying mechanisms of these diversity effects remain poorly understood. Because resource C : N : P stoichiometry (i.e. the ratios of these key elements) exhibits strong control on consumers, we supposed that stoichiometric dissimilarity of litter mixtures (i.e. the divergence in C : N : P ratios among species) improves resource complementarity to decomposers leading to faster mixture decomposition. We tested this hypothesis with: (i) a wide range of leaf litter mixtures of neotropical tree species varying in C : N : P dissimilarity, and (ii) a nutrient addition experiment (C, N and P) to create stoichiometric similarity. Litter mixtures decomposed in the field using two different types of litterbags allowing or preventing access to soil fauna. Litter mixture mass loss was higher than expected from species decomposing singly, especially in presence of soil fauna. With fauna, synergistic litter mixture effects increased with increasing stoichiometric dissimilarity of litter mixtures and this positive relationship disappeared with fertilizer addition. Our results indicate that litter stoichiometric dissimilarity drives mixture effects via the nutritional requirements of soil fauna. Incorporating ecological stoichiometry in biodiversity research allows refinement of the underlying mechanisms of how changing biodiversity affects ecosystem functioning.     

土壤动物对川西高山/亚高山森林凋落物分解的贡献

采用凋落物分解袋法,研究了土壤动物对川西高山/亚高山森林代表性植物康定柳、方枝柏、红桦和岷江冷杉凋落物在分解第一年(2011年11月-2012年10月)不同关键时期质量损失的贡献.结果表明: 在凋落物第一年的分解过程中, 不同物种凋落物的分解速率大小依次为康定柳＞红桦＞岷江冷杉＞方枝柏,且均为生长季节大于冻融季节.土壤动物对凋落物分解的贡献率(P_fau)为方枝柏(26.7%)＞岷江冷杉(18.8%)＞红桦(15.7%)＞康定柳(13.2%),其中康定柳和方枝柏的P_fau在生长季节大于冻融季节,而红桦和岷江冷杉的P_fau为冻融季节大于生长季节.冻融季节土壤动物的作用与凋落物初始C、P和N/P显著相关,而生长季节则与N、C/N、木质素、木质素/纤维素显著相关.     

三江平原草甸湿地土壤呼吸和枯落物分解的CO2释放

利用静态箱-碱液吸收法研究了三江平原草甸湿地土壤呼吸和枯落物分解的CO2释放速率,讨论了影响CO2释放的环境因素,估算了枯落物分解的CO2释放对于总释放的贡献.结果表明,生长季,小叶章沼泽化草甸和小叶章湿草甸各部分CO2释放均具有明显的时间变化特征,温度和水分是重要制约因素.两类草甸湿地的平均土壤呼吸速率分别为4.33g·m-2·d-1和6.15g·m-2·d-1,枯落物分解的CO2平均释放速率分别为1.76g·m-2·d-1和3.10g·m-2·d-1,枯落物分解的CO2释放占总释放量的31%和35%,说明在碳素由地上植物碳库转移到地下土壤碳库的过程中,湿地枯落物是一个不可忽略的碳损失源.     

Combined effects of elevated temperatures and reduced leaf litter quality on the life-history parameters of a saprophagous macroarthropod

Because soil macroinvertebrates strongly modify decomposition processes, it is important to know how their abundance will respond to global change. We investigated in laboratory microcosms, the effects of elevated temperatures and reduced leaf litter quality on the life‐history traits of a saprophagous macroarthropod (development time, growth, survival and reproduction). Millipedes (Polydesmus angustus) from an Atlantic temperate forest were reared throughout their life cycle (≥16 months) under two temperature regimes differing on average by 3.3 °C; in a factorial design, they were fed either on Atlantic leaf litter or on Mediterranean leaf litter with a higher C : N ratio; humidity was consistently high. The components of the population growth rate (r) were affected positively by the temperature rise and negatively by the switch from Atlantic to Mediterranean leaf litter. When both treatments were combined, litter effects offset temperature effects. These results show that the short‐term response of saprophagous macroarthropods to warming is positive but depends on the availability of high‐quality litter, which is difficult to predict in the global change context. In a parallel experiment, conspecific millipedes from a Mediterranean population, which have evolved for a long time in a warmer climate and on poor‐quality litter, were reared at elevated temperatures on Mediterranean leaf litter. All components of r were higher than in the Atlantic population under the same conditions. This suggests that in the longer term, macroarthropods can overcome detrimental trophic interactions. Based on our study and the literature, we conclude that for decades the positive effects of warming on saprophagous macrofauna should exceed the negative effects of changes in litter quality. The abundance of those organisms in temperate forests could increase, which is confirmed by latitudinal patterns in Europe. Studies aimed at predicting the impacts of global change on decomposition will need to consider interactions with soil macroinvertebrates.     

Decomposability of C3 and C4 grass litter sampled under different concentrations of atmospheric carbon dioxide at a natural CO2 spring

Elevated concentrations of atmospheric CO₂ can influence the relative proportions, biomass and chemical composition of plant species in an ecosystem and, thereby, the input of litter nutrients to soil. Plant growth under elevated CO₂ appears to have no consistent effect on rates of litter decomposition; decomposition can, however, differ in C3 and C4 plant material from the same CO₂ environment. We here describe the decomposability of leaf litter of two grass species – the C3 Holcus lanatus L. (Yorkshire fog) and C4 Pennisetum clandestinum Hochst. (kikuyu) - from an unfertilized, ungrazed grassland at a cold CO₂ spring in Northland, New Zealand. Decomposability was measured by net CO₂–C production from litter incubated for 56 days at 25 °C in a gley soil from the site; net mineral-N production from litter was also determined. Both litter and soils were sampled under `low' and `high' concentrations of atmospheric CO₂. Decomposition of H. lanatus litter was greater than that of P. clandestinum litter throughout the 56-day incubation. Decomposition tended to be greater in `high-CO<sub>2</sub>' than in `low-CO₂' H. lanatus litter, but lower in `high-CO<sub>2</sub>' than `low-CO₂' P. clandestinum litter; differences were, however, non-significant after 28 days. Overall, litter decomposition was greater in the `low-CO<sub>2</sub>' than `high-CO₂' soil. Differences in decomposition rates were related negatively to litter N concentrations and positively to C:N ratios, but were not predictable from lignin:total N ratios. Net mineral-N production from litter decomposition did not differ significantly in `high-CO<sub>2</sub>' and `low-CO₂' samples incubated in `low-CO<sub>2</sub>' soil; in `high-CO₂' soil some net immobilization was observed. Overall, results indicate the likely complexity of litter decomposition in the field but, nevertheless, strongly suggest that rates of decomposition will not necessarily decline in a `high-CO₂' environment.     

Climatic effects on litter decomposition from arctic tundra to tropical rainforest

Climatic effects on the decomposition rates of various litter types in different environments must be known to predict how climatic changes would affect key functions of terrestrial ecosystems, such as nutrient and carbon cycling and plant growth. We developed regression models of the climatic effects on the first‐year mass loss of Scots pine needle litter in boreal and temperate forests across Europe (34 sites), and tested the applicability of these models for other litter types in different ecosystems from arctic tundra to tropical rainforest in Canada (average three year mass loss of 11 litter types at 18 sites), the USA and Central America (four litter types at 26 sites). A temperature variable (annual mean temperature, effective temperature sum or its logarithm) combined with a summer drought indicator (precipitation minus potential evapotranspiration between May and September) explained the first‐year mass loss of the Scots pine needle litter across Europe with a higher R² value than actual evapotranspiration (0.68–0.74 vs. 0.51) and with less systematic error for any sub‐region. The model with temperature sum and the summer drought indicator appeared best suited to the other litter types and environments. It predicted the climatic effects on the decomposition rates in North and Central America with least systematic error and highest R² values (0.72–0.80). Compared with Europe, the decomposition rate was significantly less sensitive to annual mean temperature in Canada, and to changes in actual evapotranspiration in the USA and Central America. A simple model distinguishing temperature and drought effects was able to explain the majority of climatic effects on the decomposition rates of the various litter types tested in the varying environments over the large geographical areas. Actual evapotranspiration summarizing the temperature and drought effects was not as general climatic predictor of the decomposition rate.     

降水量变化对蒙古栎落叶分解过程的间接影响

分析了在4种不同降水量条件下蒙古栎叶凋落物基质质量的变化,并应用分解袋法研究其凋落物在蒙古栎次生林内的分解过程.结果表明：与对照相比,降水量减少条件下,蒙古栎叶凋落物的初始N、P、K浓度显著升高,初始木质素浓度显著降低,凋落物分解速率大,N、P、K矿化率高,N和P固持时间缩短;降水量增加情况下,其凋落物初始N浓度显著降低、木质素浓度显著升高,N、P、K矿化率低,N和P固持时间延长.4种类型叶片凋落物的质量损失过程均符合指数降解模型,分解速率可以由凋落物木质素/N来预测.相关性分析显示,木质素浓度高、N浓度低的两种凋落物的分解速率与N浓度相关性最大;而木质素浓度低、N浓度高的两种凋落物的分解速率与木质素浓度相关性最大.说明降水量的变化显著地改变了蒙古栎叶凋落物的基质质量,进而间接地改变了凋落物的分解过程.     

杉木凋落物对土壤有机碳分解及微生物生物量碳的影响

利用¹³C稳定同位素示踪技术,研究了杉木凋落物对杉木人工林表层(0～5 cm)和深层(40～45 cm)土壤有机碳分解、微生物生物量碳和可溶性碳动态的影响.结果表明: 杉木人工林中深层土壤有机碳分解速率显著低于表层土壤,但其激发效应却显著高于表层土壤.杉木凋落物添加使土壤总微生物生物量碳和源于原有土壤的微生物生物量碳均显著增加,但对土壤可溶性碳没有显著影响.深层土壤被翻到林地表层,可能加速杉木人工林土壤中碳的损失.     

Effects of elevated CO2 and nitrogen fertilization pretreatments on decomposition on tallgrass prairie leaf litter

Standing dead and green foliage litter was collected in early November 1990 from Andropogon gerardii (C₄), Sorghastrum nutans (C₄), and Poa pratensis (C₃) plants that were grown in large open-top chambers under ambient or twice ambient CO₂ and with or without nitrogen fertilization (45 kg N ha⁻¹). The litter was placed in mesh bags on the soil surface of pristine prairie adjacent to the growth treatment plots and allowed to decay under natural conditions. Litter bags were retrieved at fixed intervals and litter was analyzed for mass loss, carbon chemistry, and total Kjeldahl nitrogen and phosphorus. The results indicate that growth treatments had a relatively minor effect on the initial chemical composition of the litter and its subsequent rate of decay or chemical composition. This suggests that a large indirect effect of CO₂ on surface litter decomposition in the tallgrass prairie would not occur by way of changes in chemistry of leaf litter. However, there was a large difference in characteristics of leaf litter decomposition among the species. Poa leaf litter had a different initial chemistry and decayed more rapidly than C₄ grasses. We conclude that an indirect effect of CO₂ on decomposition and nutrient cycling could occur if CO₂ induces changes in the relative aboveground biomass of the prairie species.