Role of foliar fungal endophytes in litter decomposition among species and population origins

Litter decomposition is a key ecosystem process which returns nutrients from dead plant material to mineral forms in the soil. We examined whether systemic fungal endophytes modulate recycling of nutrients directly by altering litter decomposition. We studied litter decomposition mediated by Epichloë endophytes in litter-bag experiments. We examined direct endophyte effects on litter decomposition in wild populations and cultivars of Schedonorus phoenix and Schedonorus pratensis. In the first experiment, endophyte presence tended to increase litter decomposition rate in cultivars of the two grass species (S. phoenix and S. pratensis). However, in the second experiment plant origin had a stronger influence than endophyte symbiosis in S. phoenix. Interestingly, the initial level of alkaloids was associated positively with decomposition in S. phoenix populations. Characteristics associated with litter quality were not clearly related to either endophytes or decomposition rate. Our results suggest that endophytes can enhance litter breakdown but their role in nutrient cycling is far more complex depending on plant population origin.     

Episodic acidification affects the breakdown and invertebrate colonisation of oak litter

1. Although European streams are now recovering chemically from acidification, biological recovery is limited. One hypothesis is that continuing acid episodes restrict acid‐sensitive species in recovering locations either through direct toxicity or by affecting ecological processes. Here, we test this hypothesis by assessing the effects of episodic acid exposure on the breakdown and macroinvertebrate colonisation of oak (Quercus robur) litter. 2. Over 83 days, acid episodes of 4 days’ duration were simulated by repeatedly transplanting litter bags of contrasting mesh size between replicate acidic and circumneutral streams around Llyn Brianne (Wales, U.K.). Results were compared against controls from circumneutral streams and circumneutral transplants, while invertebrates colonising litter were compared with adjacent assemblages. 3. Breakdown was retarded significantly by repeated acid exposure in comparison with circumneutral transplants, but only in litter to which invertebrates had access. Overall breakdown was also significantly slower in fine‐mesh than in coarse‐mesh bags. 4. Plecopteran shredders were the major invertebrate colonists of litter, along with smaller numbers of grazers and predators. However, acid exposure eliminated or suppressed acid‐sensitive families, resulting in an overall composition converging on that in acid streams. 5. The rapid loss of sensitive invertebrates from acid‐exposed litter supports the hypothesis that acid episodes suppress biological recovery from acidification through direct physiological effects. However, our litter breakdown data indicate that (i) some effects of acid episodes could be mediated through litter processing; and (ii) episodic acidification could disrupt litter breakdown through effects on invertebrate composition or activity. These data suggest that delayed biological recovery from acidification can reflect a combination of direct toxic and indirect ecological effects.     

Fishpond dams affect leaf‐litter processing and associated detritivore communities along intermittent low‐order streams

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Forest plantation diversification with Leguminosae tree species: Consequences on litter decomposition and nutrient recycling

Mixed tree plantations provide greater ecosystem services than monocultures. Leguminosae tree species can be appropriate complements to achieve a sustainable soil management target. A key aspect of species trait complementarity is the litter mixture effects in the litter decomposition process. We evaluated how the mixture of poplar litter (Populus deltoides Marsh.) with Leguminosae tree species modulated the litter decomposition process and C, N and P recycling, through changes driven by the Leguminosae litter chemical traits. Under field conditions, we compared poplar litter alone (monoculture) with its 50:50 mixture with Enterolobuim contortisiliquum (Vell.) Morong., or Peltophorum dubium (Spreng.). Compared to poplar litter, its mixture with E. contortisiliquum had a 25% lower C:N ratio and a similar N:P ratio, whereas mixture with P. dubium had a 9% lower C:N and a 29% lower N:P ratios. The mixture with E. contortisiliquum showed a 64% faster decomposition rate, and 55% and 203% faster C and N release rates, respectively, compared to poplar. In contrast, in the mixture with P. dubium, there was no difference in the litter, C and N decay rates with poplar litter alone. The mixture with P. dubium had a 37% lower P retention compared to poplar, whereas P was released rather than retained in the mixture with E. contortisiliquum. The mixture with E. contortisiliquum showed a net antagonistic effect in the litter decomposition rate. However, in the mixture, poplar litter decomposed 33% faster and the E. contotrtisiliquum litter decomposed 35% slower than species alone. The C:N and N:P ratios in the litter mixture were relevant traits shaping the magnitude and direction of litter decomposition and nutrient recycling processes. The incorporation of both Leguminosae to monospecific poplar plantations could contribute to counteract P limitation in this system and to improve soil fertility and functioning.     

Aphids decelerate litter nitrogen mineralisation through changes in litter quality

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Plant diversity does not buffer drought effects on early‐stage litter mass loss rates and microbial properties

Human activities are decreasing biodiversity and changing the climate worldwide. Both global change drivers have been shown to affect ecosystem functioning, but they may also act in concert in a non‐additive way. We studied early‐stage litter mass loss rates and soil microbial properties (basal respiration and microbial biomass) during the summer season in response to plant species richness and summer drought in a large grassland biodiversity experiment, the Jena Experiment, Germany. In line with our expectations, decreasing plant diversity and summer drought decreased litter mass loss rates and soil microbial properties. In contrast to our hypotheses, however, this was only true for mass loss of standard litter (wheat straw) used in all plots, and not for plant community‐specific litter mass loss. We found no interactive effects between global change drivers, that is, drought reduced litter mass loss rates and soil microbial properties irrespective of plant diversity. High mass loss rates of plant community‐specific litter and low responsiveness to drought relative to the standard litter indicate that soil microbial communities were adapted to decomposing community‐specific plant litter material including lower susceptibility to dry conditions during summer months. Moreover, higher microbial enzymatic diversity at high plant diversity may have caused elevated mass loss of standard litter. Our results indicate that plant diversity loss and summer drought independently impede soil processes. However, soil decomposer communities may be highly adapted to decomposing plant community‐specific litter material, even in situations of environmental stress. Results of standard litter mass loss moreover suggest that decomposer communities under diverse plant communities are able to cope with a greater variety of plant inputs possibly making them less responsive to biotic changes.     

Long‐term responses of leaf litter decomposition to temperature,litter quality and litter mixing in plateau wetlands

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陆地生态系统混合凋落物分解研究进展

凋落物分解在陆地生态系统养分循环与能量流动中具有重要作用,是碳、氮及其他重要矿质养分在生态系统生命组分间循环与平衡的核心生态过程。自然生态系统中,植物群落大多具有较高的物种丰富度和多样性,其混合凋落物在分解过程中也更有可能发生养分传递、化学抑制等种间互作,形成多样化的分解生境,多样性较高的分解者类群以及复杂的级联效应分解,这些因素和过程均对研究混合凋落物分解过程、揭示其内在机制形成了极大的挑战。从构成混合凋落物物种丰富度和多样性对分解生境、分解者多样性及其营养级联效应的影响等方面,综合阐述混合凋落物对陆地生态系统凋落物分解的影响,探讨生物多样性在凋落物分解中的作用。通过综述近些年的研究发现,有超过60%的混合凋落物对其分解速率的影响存在正向或负向的效应。养分含量有差异的凋落物混合分解过程中,分解者优先利用高质量凋落物,使低质量的凋落物反而具有了较高的养分有效性,引起低质量凋落物分解加快并最终使混合凋落物整体分解速率加快;而凋落物物种丰富度对土壤动物群落总多度有轻微的影响或几乎没有影响,但是对线虫和大型土壤动物的群落组成和多样性有显著影响,并随着分解阶段呈现一定动态变化;混合凋落物改变土壤微生物生存的理化环境,为微生物提供更多丰富的分解底物和养分,优化微生物种群数量和群落结构及其分泌酶的活性,并进一步促进了混合凋落物的分解。这些基于植物-土壤-分解者系统的动态分解过程的研究,表明混合凋落物分解作用不只是经由凋落物自身质量的改变,更会通过逐级影响分解者多样性水平而进一步改变分解速率和养分释放动态,说明生物多样性确实在一定程度上调控凋落物分解及其养分释放过程。     

Evolution of litter size

Theories on the evolution of litter size among organisms were reexamined. The competition theories, including that based on the r−K-selection hypothesis, could not explain well why low-fecundity strategies have often evolved in stressful environments such as mountain streams, deep sea and the antarctic, where interspecies competition is considered to be lax. The theory, based on Itô's (1980) concept of the procurability of food by the young, was considered to have greater generality because it could explain not only the above-mentioned cases but also those where small litter size is observed in habitats with high species diversity (where interspecific competition may be keen), such as tropical rain forest. Examination of the process of selection of high-fecundity and low-fecundity genotypes also suggested that the procurability of food by the young can best explain the evolution of low-fecundity. The concept of density-induced dispersal and a distinction between density-dependent and density-independent predation pressures should be incorporated into our discussions on the evolution of reproductive rates.     

Decomposition of bracken litter

Investigations on the decomposition of bracken petioles, over a five-year period on six adjacent soil types, including moder-type humus, mull and peat, are reviewed. Changes in gross physical features, chemical composition, pH and dry weight are outlined. The succession of colonizing fungi is described and related to fungal activities.
Until the petioles were buried in the litter layer, decomposition occurred at different rates on the various sites, the rate on moder > mull > peat, but the sequence of events was similar. Large proportions of readily leached components were removed in the first few months, but 95% loss of dry matter was estimated to occur only after 11–23 years. The majority of fungi were species cosmopolitan on litter, the population becoming less specialized as decay advanced. The succession resembled those on some other woody tissues, lignin and cellulose decomposers predominating before sugar fungi. From field observations and laboratory experiments, the Basidiomycete Mycena galopus (Pers. ex Fr.) Kummer appeared to be the most active of the fungal decomposers.
Some ecological and economic implications of the decomposition of bracken litter are briefly discussed, including its effect on soil type, and advantages of bracken compared with straw as bedding for farm animals.     

Nitrogen in shoot litter,root litter and root exudates from nitrogen-fixingAlnus incana

Summary A pot experiment withAlnus incana (L.) Moench growing in sand was set up to compare the amounts of nitrogen released from plants shoot litter with that released below ground as root litter and/or root exudation. No nitrogen fixation by free-living microorganisms was found in the sand and the increased nitrogen content of the plant + soil system was therefore due to nitrogen fixation byFrankia in the alder root-nodules. Most of the nitrogen released from the plants was in the nitrogen-rich leaf and other shoot litter. Only small amounts of nitrogen were found in the drainage water from the pots and were recorded as increased nitrogen content of the sand.     

Leaf litter quality drives litter mixing effects through complementary resource use among detritivores

To comprehend the potential consequences of biodiversity loss on the leaf litter decomposition process, a better understanding of its underlying mechanisms is necessary. Here, we hypothesize that positive litter mixture effects occur via complementary resource use, when litter species complement each other in terms of resource quality for detritivores. To investigate this, monocultures and mixtures of two leaf litter species varying in quality were allowed to decompose with and without a single macro-detritivore species (the terrestrial woodlice Oniscus asellus). Resource quality of the mixture was assessed by the mean concentration, the dissimilarity in absolute and relative concentrations, and the covariance between nitrogen (N), phosphorus (P) and calcium (Ca) supply. Our results clearly show that litter mixing effects were driven by differences in their resource quality for detritivores. In particular, complementary supply of N and P was a major driver of litter mixing effects. Interestingly, litter mixing effects caused by the addition of woodlice were predominantly driven by N dissimilarity, whereas in their absence, increased P concentration was the main driver of litter mixing effects. These results show that ultimately, litter diversity effects on decomposition may be driven by complementary resource use of the whole decomposer community (i.e., microbes and macro-detritivores).     

Effects of temperature and litter type on fungal growth and decomposition of leaf litter

The dependence of fungal decomposition of leaf litter on incubation temperature and litter types used as substrata was assessed under pure culture conditions. Isolates of Xylaria sp., a major ligninolytic fungus in cool temperate forests in Japan, were used as the fungal material. Xylaria sp. is mesophilic; maximum growth and decomposition occurred at 25°C. In the temperature test, the decomposition pattern of beech leaf litter by three isolates of Xylaria sp. changed at a threshold at 25°C. Cellulolytic activity increased with temperature from 5 to 25°C, whereas above 25°C ligninolytic activity increased at the expense of cellulolytic activity, leading to suppressed overall decomposition as a result of the higher temperature. The mass loss of leaf litter caused at 20°C by an isolate of Xylaria sp. was variable among 15 litter types and was correlated negatively with acid-unhydrolyzable residue (AUR) content and positively with total carbohydrate content for the 15 litter types. The effects of temperature and litter type on the growth and decomposition of leaf litter by Xylaria sp. may have implications for changes in fungal decomposition of leaf litter that would be predicted in response to future environmental changes.     

根系在凋落物层生长对凋落叶分解及酶活性的影响

根系向凋落物层生长是森林生态系统存在的普遍现象,研究根系存在对凋落物分解的影响对理解森林生态系统的养分物质循环具有重要意义.在福建三明市楠木和格氏栲林进行1年的凋落叶分解试验,设置有根处理和无根处理(对照),研究根系生长对凋落叶分解速率、养分释放和酶活性的影响.结果表明:在分解360 d后,有根处理楠木和格氏栲凋落叶干...     

Connecting litter quality,microbial community and nitrogen transfer mechanisms in decomposing litter mixtures

Synergistic effects on decomposition in litter mixtures have been suggested to be due to the transfer of nitrogen from N‐rich to N‐poor species. However, the dominant pathway and the underlying mechanisms remain to be elucidated. We conducted an experiment to investigate and quantify the control mechanisms for nitrogen transfer between two litter species of contrasting nitrogen status (¹⁵N labeled and unlabeled Fagus sylvatica and Fraxinus excelsior) in presence and absence of micro‐arthropods. We found that ¹⁵N was predominantly transferred actively aboveground by saprotrophic fungi, rather than belowground or passively by leaching. However, litter decomposition remained unaffected by N‐dynamics and was poorly affected by micro‐arthropods, suggesting that synergistic effects in litter mixtures depend on complex environmental interrelationships. Remarkably, more ¹⁵N was transferred from N‐poor beech than N‐rich ash litter. Moreover, the low transfer of ¹⁵N from ash litter was insensitive to destination species whereas the transfer of ¹⁵N from labeled beech litter to unlabeled beech was significantly greater than the amount of ¹⁵N transferred to unlabeled ash suggesting that processes of nitrogen transfer fundamentally differ between litter species of different nitrogen status. Microbial analyses suggest that nitrogen of N‐rich litter is entirely controlled by bacteria that hamper nitrogen capture of microbes in the environment supporting the source‐theory. In contrast, nitrogen of N‐poor fungal dominated litter is less protected and transferable depending on the nitrogen status and the transfer capacity of the microbial community of the co‐occurring litter species supporting the gradient‐theory. Thus, our results challenge the traditional view regarding the role of N‐rich litter in decomposing litter mixtures. We rather suggest that N‐rich litter is only a poor nitrogen source, whereas N‐poor litter, can act as an important nitrogen source in litter mixtures. Consequently both absolute and relative differences in initial litter C/N ratios of co‐occurring litter species need to be considered for understanding nitrogen dynamics in decomposing litter mixtures.     

Interacting effects of leaf litter species and macrofauna on decomposition in different litter environments

The leaf litter environment (single species versus mixed species), and interactions between litter diversity and macrofauna are thought to be important in influencing decomposition rates. However, the role of soil macrofauna in the breakdown of different species of leaf litter is poorly understood. In this study we examine the multiple biotic controls of decomposition – litter quality, soil macrofauna and litter environment and their interactions. The influence of soil macrofauna and litter environment on the decomposition of six deciduous tree species (Fraxinus excelsior L., Acer pseudoplatanus L., Acer campestre L., Corylus avellana L., Quercus robur L., Fagus sylvatica L.) was investigated in a temperate forest, Wytham Woods, Southern England. We used litterbags that selectively excluded macrofauna to assess the relative importance of macrofauna versus microbial, micro and mesofauna decomposition, and placed single species bags in either conspecific single species or mixed species litter environments. The study was designed to separate plant species composition effects on litter decomposition rates, allowing us to evaluate whether mixed species litter environments affect decomposition rates compared to single species litter environments, and if so whether the effects vary among litter species, over time, and with regard to the presence of soil macrofauna. All species had faster rates of decomposition when macrofauna were present, with 22–41% of the total mass loss attributed to macrofauna. Macrofauna were most important for easily decomposable species as soon as the leaves were placed on the ground, but were most important for recalcitrant species after nine months in the field. The mass loss rates did not differ between mixed and single species litter environments, indicating that observed differences between single species and mixed species litterbags in previous field studies are due to the direct contact of neighbouring species inside the litterbag rather than the litter environment in which they are placed.     

Analysis of litter size and average litter weight in pigs using a recursive model

An analysis of litter size and average piglet weight at birth in Landrace and Yorkshire using a standard two-trait mixed model (SMM) and a recursive mixed model (RMM) is presented. The RMM establishes a one-way link from litter size to average piglet weight. It is shown that there is a one-to-one correspondence between the parameters of SMM and RMM and that they generate equivalent likelihoods. As parameterized in this work, the RMM tests for the presence of a recursive relationship between additive genetic values, permanent environmental effects, and specific environmental effects of litter size, on average piglet weight. The equivalent standard mixed model tests whether or not the covariance matrices of the random effects have a diagonal structure. In Landrace, posterior predictive model checking supports a model without any form of recursion or, alternatively, a SMM with diagonal covariance matrices of the three random effects. In Yorkshire, the same criterion favors a model with recursion at the level of specific environmental effects only, or, in terms of the SMM, the association between traits is shown to be exclusively due to an environmental (negative) correlation. It is argued that the choice between a SMM or a RMM should be guided by the availability of software, by ease of interpretation, or by the need to test a particular theory or hypothesis that may best be formulated under one parameterization and not the other.