Litter Quality and Soil Microorganisms Mediate Reduced Litter Decomposition Following Understory Vegetation Removal in Forest Ecosystems

Litter decomposition was retarded by understory removal but was unaffected by thinning in a Chinese fir [Cunninghamia lanceolata (Lamb.) Hook] plantation

Despite the importance of understory vegetation for ecosystem processes and functions, its contribution to the dynamics of phosphorus (P) fractions in decomposing litter has rarely been studied in subtropical plantations. Herein, by conducting an understory removal (UR) experiment in a moso bamboo (Phyllostachys edulis) plantation of South China, we assessed the effect of understory species on leaf litter mass loss and P fractions (inorganic P, sugar P, nucleic P, and residual P) of moso bamboo over a 12‐month incubation period. We also examined the effect of UR on soil physicochemical properties, as well as fungal and bacterial phospholipid fatty acids (PLFAs). Our results showed that UR reduced litter decomposition rate (6.0–12.4%), but exhibited no effect on total P remaining in decomposing litter. Furthermore, UR produced variable effects on the amounts of P fractions in decomposing litter. Despite no changes were observed in the amount of inorganic and sugar P fractions, UR accelerated nucleic P release (3.0–6.6%), and residual P immobilization (9.2–27.9%) in decomposing litter. Moreover, UR increased fungal PLFAs (8.0–44.9%) but decreased bacterial PLFAs (18.7–36.8%) in the soil, leading to an increase in fungal to bacterial PLFAs ratio (34.5–80.9%). The correlation analysis showed that accelerated litter nucleic P release after UR was mainly attributed to the increases in soil fungal PLFAs, while decelerated litter residual P release was mainly resulted from the increased soil fungal PLFAs and decreased soil bacterial PLFAs. These results indicate that understory vegetation can mediate the dynamics of litter P fractions via its effect on soil microbial composition in subtropical plantations and suggest that understory vegetation is crucial to maintain soil P availability in moso bamboo plantations.

Improvement cutting or harvesting can change the coverage of understory vegetation, which can significantly influence the litter decomposition process in plantations. However, difference in potential non-additive mass loss in response to understory vegetation changes is poorly studied. A field litterbag experiment involving various litter types and treatments with no understory vegetation removal, shrub removal, herb removal and whole-understory vegetation removal was conducted to examine non-additive mass loss. During approximately 2 years of decomposition, the decomposition rate of shrub and herb components was accelerated in the mixed litter with full understory vegetation. There was significant non-additive mass loss during decomposition in the plots with trees, shrubs and herbs, while the incidence of non-additive mass loss was lower in the plots with understory vegetation removal. Statistical analysis revealed a significant difference between the expected mass loss calculated with the data from the corresponding decomposition plots and that calculated with the data from the plots with whole-understory vegetation removal. Our results show that understory vegetation removal can inhibit litter decomposition in Masson pine plantation ecosystems in subtropical China. We highlight that non-additive litter decomposition should be assessed on the basis of litter species composition and decomposition microenvironments in situ.

高山森林林窗对凋落叶分解的影响

Summary Increasing atmospheric CO2 concentrations alongside more frequent and severe droughts are key global change factors impacting litter decomposition, and global carbon and nitrogen cycles. Yet, we have a poor understanding of how these perturbations impact interactions between litter chemistry and decomposer food webs. We tested how drought and elevated CO2 concentrations modify litter decomposition via litter chemistry and the decomposer communities using two separate, long-term manipulative drought or elevated CO2 field experiments in mature oak woodlands. Litter bags were deployed in a reciprocal transplant design whereby control and treated litter was incubated in control and treated plots. We measured litter mass loss, quality and stoichiometry, moisture content, and microbial and animal community composition. Elevated CO2 and drought affected litter stoichiometry and quality which in turn affected decomposition: litter in droughted plots decomposing slower than controls, while litter from elevated CO2 plots decomposed slower over the first three harvests. In addition, we found that litter mass loss and decomposers were related both directly and indirectly to litter C:N. Drought and elevated CO2 have distinct pathways of influence on litter quality, decomposers and decomposition which challenges our ability to predict how combinations of factors influence litter decomposition. Plain Language Summary Global change factors like drought and elevated atmospheric CO2 concentrations can impact litter decompositions, an important ecosystem process, via changes to litter properties and the decomposer community. Examining links between litter properties, decomposers and decomposition is therefore critical to understand how both drought and elevated CO2 will affect nutrient release and cycling of belowground environments.

Litter decomposition serves an important role in maintaining nitrogen (N) availability within forest ecosystems. However, the interactive effects of exogenous N, drought, and litter quality and mixing on N immobilization during decomposition remain unclear. The aim of this study was to assess the effects of litter quality, reduced precipitation, N addition, and their interactions on litter mass loss and N immobilization. This field study analyzed the effects of N addition and decreased precipitation on the decomposition rates and associated N immobilization of four types of litter: Quercus mongolica (QM), Tilia amurensis (TA), Pinus koraiensis (PK), and a mixture (MIX) of all three. The chemical quality of the MIX was prepared in a 4:3:3 (mass) ratio of PK, TA, and QM litters. Litterbags were placed in an N addition and precipitation manipulation forest field and collected after 92, 154, 365, 457, and 874 days. Decomposing litter residues were characterized for mass loss and N content to assess N immobilization. The addition of N had no significant effect on litter decomposition under both precipitation conditions, but a reduction in precipitation significantly depressed litter decomposition. The increases in N immobilization with N addition depended on the litter type and decomposition period. Precipitation reduction had significant effects on N immobilization and enhanced the magnitude and duration of N immobilization in decomposing litter, and both of which can be increased by N addition. The results indicate that the litter species is the major regulator that controls mass loss and N immobilization. Furthermore, the MIX treatment did not show non-additive effects on mass loss but did exhibit some weak synergistic effects on N immobilization. Our results suggest that decomposing litters could help to sequester N depending on the litter identity and water regime in temperate forest ecosystems.

Clarifying the functional consequences of intraspecific trait variability in response to interacting trophic levels would provide a significant improvement in our understanding of above‐ground–below‐ground linkages. In particular, the effects of grazing on plant traits may translate into altered litter quality, with potentially important consequences for litter‐feeding decomposers. Plant and litter variability in response to grazing is expected to depend on soil fertility levels, with tolerance and defensive strategies more commonly expressed on fertile and poorer soils, respectively. However, how grazing and fertility interactively alter litter quality and palatability to detritivores has not been explored yet. We conducted a cafeteria experiment with three common millipede (Diplopoda) species feeding on leaf litter from two plant species, the grass Bromopsis erecta and the forb Potentilla verna. Each millipede was offered a binary choice between litter types produced by the same plant species, but sampled in plots with distinct herbivory and fertilization status: litter originating from grazed areas or from 1‐year sheep exclosures, both in native areas and in adjacent paddocks that received chemical N and P fertilization, as well as litter from a 25‐year sheep exclosures in the native area. We found that fertilization and herbivore exclusion interactively affected Bromopsis litter quality and palatability, whereas Potentilla was much less affected. Bromopsis litter palatability was not affected by grazing when litter was collected in native plots, except for the long‐term exclosure which led to low palatability. In contrast, and in line with our expectations, herbivory was associated with much higher palatability in fertilized plots. The changes in palatability were associated with important alterations of litter quality. Overall, our study demonstrates that intraspecific variation in litter can have profound consequences for soil functioning. It emphasizes the role of grazing as a key, but plant species‐specific factor controlling litter intraspecific variability, and its complex interaction with soil fertility level. Moreover, our results advocate for a better understanding of the response of the different organisms involved in the decomposition process, in particular litter‐feeding macro‐detritivores. We encourage future studies aiming at disentangling the various and complex relationships between above‐ground processes such as herbivory and soil functioning.

Summary 1. Soil micro-organisms play important roles in ecosystems and respond quickly to environmental changes. We examined how understory removal and tree girdling influence the composition of soil microbial community and the litter decomposition in two subtropical plantations. 2. Phospholipid fatty acids (PLFAs) analysis was used to characterize soil microbial community. Redundancy analysis and principal response curves (PRC) were used to investigate the relationships between soil microbial community and environmental factors. 3. Understory removal significantly reduced the amount of fungal PLFAs, the ratio of fungal to bacterial PLFAs, and the litter decomposition but did not affect bacterial PLFAs and total PLFAs. In contrast, tree girdling did not affect the soil microbial characteristics. The changes in soil microbial community caused by understory removal were mainly attributed to the indirect effects such as increased soil temperature and soil NO3 ) -N availability. In addition, PRC analysis showed that the relative abundance of most PLFAs increased in response to understory removal in the 2-year-old plantation but decreased in the 24-year-old plantation. 4. We propose that understory plants are important components in subtropical forest ecosystems, and play different roles in maintaining soil microbial community and driving litter decomposition processes in young vs. old plantations. The functions of understory plants should be considered in forest management and restoration. The negligible effect of tree girdling on the soil micro-organisms can be attributed to the resprouting trait and mycorrhizal interactions of Eucalyptus.

Nutrient enrichment and the subsequent chemical changes in dystrophic savanna soils may alter plant richness and nutrient use efficiency, thereby affecting leaf litter chemistry and decomposition. However, the role of soil mesofauna in litter decomposition in savanna ecosystems under nutrient enrichment is not well understood. In soils from a long‐term fertilisation experiment, we evaluated the decay of leaf litter incubated in fine‐ and coarse‐mesh bags over a year to assess the role of soil mesofauna in litter decomposition in the central Brazilian savannas. Experimental plots were established in a woodland savanna and consisted of nitrogen (N), phosphorus (P), N plus P and lime additions and untreated control (three replicates of each). We evaluated the effect of fertilisation and liming on litter decomposition rates (total, mesofauna‐mediated and microbe‐mediated) and their relationship with the initial litter quality (N, C, lignin, cellulose and polyphenol content). Litter decomposition rates were significantly higher in coarse‐mesh bags in the N plus P treatment compared with the control, which was explained by mesofauna‐mediated decomposition but not by the initial litter quality. Litter mass losses in fine‐mesh bags were significantly higher in the initial months of the experiment in the N treatment and in the intermediate months in the N plus P treatment compared with the control. This result could be explained by the initial litter N content, although it has not reflected in significantly higher microbe‐mediated decomposition rates. The litter mass losses in coarse‐mesh bags in the N plus P treatment were significantly higher only in the final months, indicating that microorganisms and mesofauna differentially affect decomposition over time. Our findings suggest that combined N and P addition could alter soil organic matter dynamics in Brazilian savannas by increasing soil mesofauna‐mediated litter decomposition rates.

BackgroundThe prevalence of understory removal and anthropogenic nitrogen (N) deposition has significantly altered the ecological processes of forest ecosystems at both regional and global scales. However, it remains a pressing challenge to understand how N deposition and understory removal affect leaf nutrient dynamics, nutrient resorption, litter decomposition, and their linkages for better managing forest ecosystems under nutrient imbalances induced by N enrichment. To address this research gap, a field manipulation experiment was carried out in a subtropical Cunninghamia lanceolata plantation with four treatments including: control (CK), canopy N addition (CN), understory removal (UR), and canopy N addition plus understory removal (CN × UR). Green and senesced leaf N and phosphorus (P) concentrations, N and P resorption efficiencies, litter decomposition, and their correlations were measured.ResultsThe results revealed that the average N concentrations of green early and late leaves in UR were increased by 6.61 and 18.89% compared to CK. UR had the highest whereas CN had the lowest P concentrations in green leaves across the two sampling seasons. Following this, UR, leaf type, season, and their interactions significantly affected leaf N, P, and N:P (P < 0.05). The highest leaf N resorption (32.68%) and P resorption efficiencies (63.96%) were recorded in UR. Litter decomposition was significantly retarded in UR (P < 0.01) relative to CN. The regression analysis demonstrated that leaf nutrient status was significantly interconnected with leaf nutrient resorption efficiencies. In addition, leaf nutrient dynamics were strongly correlated with litter nutrients, indicating that both were coupled.ConclusionThese findings can deepen our knowledge of biogeochemical cycling and reveal contrasting nutrient-acquisition strategies on N and P limitation in response to UR and CN. Considering the P limitation, it is important to note that P was resorbed more efficiently, illustrating a remarkable nutrient preservation approach for nutrient-limitations. Resorption may be a crucial mechanism for keeping nutrients in these forests, so better understory management practices are required to prevent reliance on external nutrient pools. Overall, this study sheds meaningful insights into the ability of forest adaptation in response to global climatic change.

Subtropical plantations have been undergoing understory removal and increased plant litter inputs due to global change and anthropogenic activities. However, how understory removal and increased litter inputs affect litter decomposition is still unclear in these plantations. Using a field manipulation experiment, we assessed the effects of understory removal and litter addition on decomposition rates of leaf litter, twig litter, and mixed leaf and twig litter over 16 months of incubation in a subtropical Chinese fir (Cunninghamia lanceolata) plantation in southern China. We also examined the changes in soil chemical properties and fungal and bacterial phospholipid fatty acids (PLFAs) after understory removal and litter addition. Litter decomposition rates were inhibited by understory removal due to the decreased fungal PLFAs and the associated declines in the ratio of fungal to bacterial biomass. Litter addition also retarded litter decomposition rates, despite the increases in soil NH4+‐N concentration and fungal PLFAs. However, both litter decomposition rates and soil microbial PLFAs remained unchanged in the presence of both understory removal and litter addition. In all treatments, mixed leaf and twig litter produced additive effects on litter decomposition rates. These results suggest that understory removal and litter addition interact to affect litter decomposition dynamics and highlight that litter mixture decomposition rates could be easily predicted from component litter in subtropical Chinese fir plantations of southern China.

Understory vegetation removal significantly affected soil biogeochemical properties in forest ecosystems

Differences among plant genotypes can influence ecosystem functioning such as the rate of litter decomposition. Little is known, however, of the strength of genotypic links between litter quality, microbial abundance and litter decomposition within plant populations, or the likelihood that these processes are driven by natural selection. We used 19 Betula pendula genotypes randomly selected from a local population in south-eastern Finland to establish a long-term, 35-month litter decomposition trial on forest ground. We analysed the effect of litter quality (N, phenolics and triterpenoids) of senescent leaves and decomposed litter on microbial abundance and litter mass loss. We found that while litter quality and mass loss both had significant genotypic variation, the genotypic variation among silver birch trees in the quantity of bacterial and fungal DNA was marginal. In addition, although the quantity of bacterial DNA at individual tree level was negatively associated with most secondary metabolites of litter and positively with litter N, litter chemistry was not genotypically linked to litter mass loss. Contrary to our expectations, these results suggest that natural selection may have limited influence on overall microbial DNA and litter decomposition rate in B. pendula populations by reworking the genetically controlled foliage chemistry of these populations.

Soil microorganisms play key roles in ecosystems and respond quickly to environmental changes. Liming and/or understory removal are important forest management practices and have been widely applied to planted forests in humid subtropical and tropical regions of the world. However, few studies have explored the impacts of lime application, understory removal, and their interactive effects on soil microbial communities. We conducted a lime application experiment combined with understory removal in a subtropical Eucalyptus L’Hér. plantation. Responses of soil microbial communities (indicated by phospholipid fatty acids, PLFAs), soil physico-chemical properties, and litter decomposition rate to lime and/or understory removal were measured. Lime application significantly decreased both fungal and bacterial PLFAs, causing declines in total PLFAs. Understory removal reduced the fungal PLFAs but had no effect on the bacterial PLFAs, leading to decreases in the total PLFAs and in the ratio of fungal to bacterial PLFAs. No interaction between lime application and understory removal on soil microbial community compositions was observed. Changes in soil microbial communities caused by lime application were mainly attributed to increases in soil pH and NO3–-N contents, while changes caused by understory removal were mainly due to the indirect effects on soil microclimate and the decreased soil dissolved carbon contents. Furthermore, both lime application and understory removal significantly reduced the litter decomposition rates, which indicates the lime application and understory removal may impact the microbe-mediated soil ecological process. Our results suggest that lime application may not be suitable for the management of subtropical Eucalyptus plantations. Likewise, understory vegetation helps to maintain soil microbial communities and litter decomposition rate; it should not be removed from Eucalyptus plantations.

Litter inputs are closely related to both forest productivity and nutrient cycling under climate change and local management. This study investigated the effect of litter inputs on litter decomposition, changes in litter chemistry and nitrogen (N) dynamics during eucalyptus leaf litter decomposition. Two parallel in situ litter decomposition experiments were conducted at two sites with high-quality (HQ) and low-quality (LQ) litters in a eucalyptus-dominated forest of southeast Queensland, Australia. At each site, leaf litters with either a single (SL) or double mass load (DL) of litter inputs were decomposed for 15 months. Litter mass loss, chemical composition and N content of decomposing litters were measured seasonally during the decomposition period. The chemical composition of the collected litters was determined by solid-state 13C nuclear magnetic resonance (NMR) spectroscopy. The HQ litters decomposed faster than the LQ litter, with a decomposition constant of 0.53 and 0.33 y−1 at the HQ and LQ site, respectively. Litter addition rates had no effect on litter decomposition, changes in chemical composition and N content during decomposition regardless of differences in initial litter quality. The HQ and LQ litters showed the same pattern of chemical changes during decomposition, with an increase in alkyl C and a decrease in di-O-alkyl C and aryl C. The relative intensity of O-aryl C and carboxyl C converged, while the relative intensity of di-O-alkyl C and δ15N diverged as the decomposition progressed. N immobilization during decomposition depended on litter quality, with N consistently immobilized in LQ litters over the whole decomposition period. In subtropical eucalyptus-dominated forests, the dynamics of organic C and N during litter decomposition were resistant to the increased inputs of aboveground litters. Litter chemistry of different initial qualities converged at the early stages of decomposition, and the implications of chemical convergence on the formation and stabilization of soil organic matter need to be assessed in the future.