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We also expected that 3 variations in microbial biomass and ergosterol concentration in litter are closely linked to changes in the litter decomposition processes. Overall, the study aims at improving the understanding of the mechanisms contributing to the accumulation of dead organic material in high altitude tropical rainforests. The study area was in southern Ecuador on the eastern slope of the Andes. Within this area we established three sites along an altitudinal gradient.

With 8—10 humid months per year the region has a semihumid climate. The mean annual air temperature is negatively related to altitude being Soil pH is similarly related to altitude being 3. Mean soil moisture in the organic layer increases with increasing altitude being 9. Biotic conditions also change along the altitudinal gradient. Mean tree height decreases with altitude being The biomass of fine roots has the same pattern with respective values of 2.

Nylon bags litterbags, 4 mm mesh were used to investigate the decomposition and microbial colonization of leaves and roots, that is, two types of litter materials. The mesh size allows access to the litter by the dominant decomposer mesofauna but minimizes the loss of litter due to handling.

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Root litter was obtained by digging up the upper 0. The roots were then cleaned of adhering organic matter and soil by rinsing them gently with tap water. The amount of leaf litter of each species and the quantity of roots from each size class placed into the litterbags of the three litter species was based on the amounts present at each study site. Leaves of the three plant families or roots of the three root size classes were mixed according to their relative abundances at each altitude.

There were thus litterbags in total. Scheme of collection of litter materials origins and placement of litterbags in the field altitudes. After retrieval, the remaining leaf and root litter were cleaned by removing roots that had grown into the litterbags.

Plant Litter

For these measurements, material from each litterbag was homogenized by cutting it into pieces of about 0. Moist samples equivalent to 0. Ergosterol was extracted from 0. Each 0. The dried extract was collected in 0. We focused on the variation in the amount and concentration of C within the litter material in order to be able to link decomposition processes closely to energetic processes.

Therefore, the amounts of C remaining C R in the litterbags at the sampling dates n were expressed as percentages of the initial amount of C placed into the litterbags C 0. Similarly, changes in the amount of N remaining N R were expressed as percentage of the initial amount of N placed in the litterbags N 0 , according to the following formulas:.

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Block was excluded from the analysis as there were no significant block effects. Before the analyses, data were inspected for homogeneity of variance and normal distribution. Means presented in the Results are based on nontransformed data. Litter type had the strongest influence on the ergosterol concentration, while C R and C mic varied most strongly with sampling date. C R and qO 2 were not affected by the origin of the litter under main effects at any sampling date.

Only N R and N C significantly varied with litter origin at all of the sampling dates. The following results therefore focus on the effects of altitude, sampling date, and litter type. Significant effects are given in bold. In contrast, much more remained at the two highest sites, Within the first year, C R decreased to a similar extent in leaf and root litter, with the reductions averaging In leaf litter, the reduction was 5.

C R in litter from Bomb did not vary significantly between leaf and root litter. The initial C C of the litter placed in the litterbags was generally high but differed between leaf litter The pattern differed in subsequent years. The C C in litter from Bomb and Caja varied similarly. It increased by 1. In the third year, the increase was 1. In the fourth year, C C decreased in both places by an average of 1. From 24 to 36, it decreased by 1. The decline continued in subsequent years at an average of Root litter lost only At higher altitudes, the decrease was also at a maximum in litter from Bomb with N R declined most strongly in leaf litter from Bomb with Initial N C of the litter materials in the litterbags was generally low but differed among sites for leaf litter 1.

In the other treatments, the increase in N C varied between 0. Overall, N C decreased with increasing altitude and was higher in leaf litter than in root litter. Litter from Bomb, which had the highest initial N C , had the lowest increase irrespective of altitude. In contrast, in litter from Caja the increase in N C in root litter 0. In contrast to this view, we showed that altitude affects decomposition by modifying microenvironmental conditions rather than the quality of the litter, that is, litter origin, which only slightly influenced the amount of C remaining C R in the early phase of decomposition.

Our results support this view, with a rapid decline in the amount of litter C within the first year of decomposition irrespective of altitude. Berg described that in the later phases of decomposition the rate of mass and C loss slows down and is dominated by the degradation of the remaining recalcitrant litter compounds such as lignin. This resumption of C loss indicates a third phase of decomposition which probably is associated with slow degradation of recalcitrant litter compounds such as lignin and a shift in the decomposer community toward lignin decomposing microorganisms.

The long delay of this resumption of decomposition suggests that the late microbial community, degrading recalcitrant litter compounds, only establishes through facilitation by the activity of the early colonisers and the degradation of labile litter compounds. Macrofauna density, therefore, cannot explain the difference in decomposition rates between the study sites. We showed in our study that decomposition patterns were similar at the two higher altitudes although temperature, precipitation, soil moisture, and soil pH varied between each of the three study sites.

Further, our study shows that precipitation and soil pH also are of little importance. We presume that the different forest floor types at the different altitudes contribute to the different decomposition dynamics. The lack of N transfer into the litter may cause a feedback loop, that is, the accumulation of organic material and the formation of thick F layers further inhibiting litter decomposition. We presume that this is because root litter contains higher concentrations of lignin and lower concentrations of N than leaf litter, resulting in a greater accumulation of recalcitrant litter compounds in the later phases of decomposition.

The initial N C of both leaf and root litter materials was generally low and increased during exposure at all the altitudes and in both litter types with the changes in N C varying with the initial N C. In litter material with initially high N C , the increase with time was less pronounced than in litter material with low N C , suggesting that N C values converge with time. This contrasts decomposition of litter materials low in N in temperate and boreal forest ecosystems which typically accumulate N for longer periods of time Berg, The reduction in N R at later stages of litter decay also varied with the initial N C.

In litter material with low initial N C , the increase in N C was stronger than in litter material with high initial N C. This suggests that plant roots at the study site are unable to obtain a sufficient amount of N from decomposing litter. Plants presumably rely on mycorrhizal fungi improving N capture by growing into leaf and root litter material. This linkage likewise was evident throughout the decomposition process in our study.

This again supports our conclusion that litter resources are of low quality and difficult to decompose due to low N and high concentrations of recalcitrant compounds. This again supports our conclusion that net N mobilization from the litter was due to trophic interactions between saprotrophic microorganisms, microbial grazers and VA mycorrhizal fungi see above.

High qO 2 during this phase see Fig. Franca Marian involved in investigation, formal analysis, and writing the original draft. Dorothee Sandmann and Valentyna Krashevska involved in investigation; formal analysis; and writing, review, and editing of the manuscript. Mark Maraun and Stefan Scheu involved in conceptualization; methodology; writing, review, and editing of the manuscript; supervision; and funding acquisition. Measurements of C and N were performed in the laboratory of Prof.

Valentyna Krashevska 1 J. Mark Maraun 1 J. Stefan Scheu 1 J. Author information Article notes Copyright and License information Disclaimer. Franca Marian, Email: ed. Corresponding author. Email: ed. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.


This article has been cited by other articles in PMC. Associated Data Supplementary Materials. Keywords: altitudinal gradient, Ecuador, litter quality, litter type, litterbag, microbial biomass. Study site The study area was in southern Ecuador on the eastern slope of the Andes. Experimental setup Nylon bags litterbags, 4 mm mesh were used to investigate the decomposition and microbial colonization of leaves and roots, that is, two types of litter materials.

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Figure 1. Analytical procedures After retrieval, the remaining leaf and root litter were cleaned by removing roots that had grown into the litterbags. Calculations and statistical analysis We focused on the variation in the amount and concentration of C within the litter material in order to be able to link decomposition processes closely to energetic processes. Figure 2.

Figure 3. Figure 4. Figure 5.


Climate, leaf litter chemistry and leaf litter decomposition in terrestrial ecosystems: A triangular relationship. Oikos , 79 3 , — Nature Geoscience , 3 5 , — A physiological method for the quantitative measurement of microbial biomass in soils. Soil Biology and Biochemistry , 10 3 , — Nature , , — Ecology , 96 3 , — Global centers of vascular plant diversity. Nova Acta Leopoldina , 92 , 61— Soil Biology and Biochemistry , 29 7 , — Decomposition patterns for foliar litter — A theory for influencing factors. Soil Biology and Biochemistry , 78 , — Plant litter: Decomposition, humus formation, carbon sequestration , 2nd ed.

Berlin, DE: Springer. Adaptive responses of soil microbial communities under experimental acid stress in controlled laboratory studies.

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  • Applied Soil Ecology , 11 2—3 , — Temperature sensitivity and enzymatic mechanisms of soil organic matter decomposition along an altitudinal gradient on Mount Kilimanjaro. Scientific Reports , 6 1 , Protozoa and plant growth: The microbial loop in soil revisited. New Phytologist , 3 , — Climatic conditions and tropical montane forest productivity: The fog has not lifted yet. Ecology , 79 1 , 3—9. Litter mixture effects on decomposition in tropical montane rainforests vary strongly with time and turn negative at later stages of decay.

    Soil Biology and Biochemistry , 77 , — Litter quality versus soil microbial community controls over decomposition: A quantitative analysis. Oecologia , 1 , — The microbial loop concept as used in terrestrial soil ecological studies. Microbial Ecology , 28 2 , — Soil Biology and Biochemistry , 34 1 , 69— Sensitivity of tropical carbon to climate change constrained by carbon dioxide variability.

    Global Change Biology , 15 5 , — Temperature sensitivity of soil carbon decomposition and feedbacks to climate change. Soil carbon stocks vary predictably with altitude in tropical forests: Implications for soil carbon storage. Geoderma , — , 59— Ergosterol and microbial biomass relationship in soil. Biology and Fertility of Soils , 22 4 , — Invasion of a deciduous forest by earthworms: Changes in soil chemistry, microflora, microarthropods and vegetation.

    Decomposition, Humus Formation, Carbon Sequestration

    Soil Biology and Biochemistry , 39 5 , — Similar response of labile and resistant soil organic matter pools to changes in temperature. Nature , , 57— Litter quality and the temperature sensitivity of decomposition. Ecology , 86 2 , — Global Change Biology , 6 7 , — Evidence that decomposition rates of organic carbon in mineral soil do not vary with temperature.

    Net primary productivity allocation and cycling of carbon along a tropical forest elevational transect in the Peruvian Andes. Global Change Biology , 16 12 , — Northern peatlands: Role in the carbon cycle and probable responses to climatic warming. Ecological Applications , 1 2 , — Plant and Soil , 1—2 , — Graffenrieda emarginata Melastomataceae forms mycorrhizas with Glomeromycota and with a member of the Hymenoscyphus ericae aggregate in the organic soil of a neotropical mountain rain forest.

    Canadian Journal of Botany , 82 3 , — Neotropical plant diversity. Nature , , 21— Soil microarthropod contribution to decomposition dynamics: Tropical—temparate comparison of a singel substrate.


    Plant Litter: Decomposition, Humus Formation, Carbon Sequestration by Björn Berg

    Ecology , 80 6 , — Diversity and composition of Arctiidae moth ensembles along a successional gradient in the Ecuadorian Andes. Diversity and Distributions , 11 5 , — An arbuscular mycorrhizal fungus accelerates decomposition and acquires nitrogen directly from organic material. Are microorganisms more effective than plants at competing for nitrogen? Trends in Plant Science , 5 7 , — Biotropica , 42 2 , — The authors can be congratulated for their impressive efforts in publishing this highly professional volume.

    Berg during the last two decades. New and Traditional Analytical Techniques. JavaScript is currently disabled, this site works much better if you enable JavaScript in your browser. Life Sciences Ecology. Free Preview. Third edition, fully revised and updated with much new material New information emphasizes on three fields: the effects of manganese on decomposition and possibly on carbon sequestration, new results on decomposition dynamics, and the new analytical technique using 13 C-NMR In-depth approach to the results of long-term studies see more benefits.

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