Litter decomposition and faunal activity in Mediterranean forest soils: effects of N content and the moss layer

Accumulation of soil carbon is mainly controlled by the balance between litter production and litter decomposition. Usually In Mediterranean forests there are contrasting conditions in the distribution of faunal activity and the moss layer that may have different effects on litter decomposition. Decomposition and faunal activity were studied by exposing litter of contrasting quality (Pinus halepensis Mill. and Quercus ilex L.) for 3.5 yr in three Mediterranean pine forests of the eastern Iberian Peninsula. The effects of mosses on decomposition and on faunal activity were studied by exposing P. halepensis litter either on moss patches or directly on the forest floor. Faecal pellet production was used as an indication of faunal activity. Water availability or soil characteristics seem to limit faunal activities in the drier sites. Faecal pellets were not found during the first stages of decomposition and in all sites they appeared when about a 30% of the initial litter had decomposed. Under wet conditions faecal pellet production was very high and a mass balance suggested that soil faunal activity may result in a net flow of organic matter from the lower organic horizons to the surface Oi horizon. Mosses slightly increased mass loss of pine litter probably as a consequence of high potentially mineralizable nitrogen in the Oa horizon of moss patches and also, perhaps, as a consequence of the higher moisture content measured in the Oi horizon needles sampled among the mosses. In contrast, moss patches reduced faunal activity. The effect of litter quality on mass loss was not always significant, suggesting an interaction between litter quality and site conditions. During the first stages of decomposition there was N immobilisation in P. halepensis litter (poorer in N) and N release from Q. ilex litter (richer in N). In conclusion, in these forests soil microclimate and/or N availability appear to be more important controlling litter decomposition than the distribution of faunal activity.     

Immobilization of avian excreta-derived nutrients and reduced lignin decomposition in needle and twig litter in a temperate coniferous forest

The possible effects of excreta of the Great Cormorant Phalacrocorax carbo on decomposition processes and dynamics of nutrients (N, P, Ca, K, Mg) and organic chemical components (lignin, total carbohydrates) were investigated in a temperate evergreen coniferous forest near Lake Biwa in central Japan. Two-year decomposition processes of needles and twigs of Chamaecyparis obtusa were examined at two sites, control site never colonized by the cormorants (site C) and colonizing site (site 2). Mass loss was faster in needles than in twigs. Mass loss of these litter types was faster at site C than at site 2, which was ascribed to the decreased mass loss rate of acid-insoluble ‘lignin’ at site 2. Net immobilization of N, P, and Ca occurred in needles and twigs at site 2; whereas at site C, mass of these elements decreased without immobilization during decomposition. Duration of immobilization phase of these nutrients at site 2 was estimated to be 1.6 to 2.5 years in needles and 19.6 to 23.5 years in twigs. Immobilization potential (maximum amount of exogenous nutrient immobilized per gram initial material) was similar between needles and twigs for N and Ca but was about 10 times higher in twigs than in needles for P. δ¹³C in needles was relatively constant during the first year and then increased during the second year, whereas δ¹³C in twigs was variable during decomposition. Acid-insoluble fraction was depleted in ¹³C compared to whole needles (1.6-2.1‰) and twigs (2.0-2.5‰). δ¹⁵N of needles and twigs and their acid-insoluble fractions approached to δ¹⁵N of excreta during decomposition at site 2. This result demonstrated the immobilization of excreta-derived N into litter due to the formation of acid-insoluble lignin-like substances complexed with excreta-derived N. No immobilization occurred in K and Mg and their mass decreased during decomposition at both sites. Based on these results of nutrient immobilization during decomposition and on the data of litter fall and excreta amount at site 2, we tentatively calculated stand-level immobilization potential of litter fall and its contribution to total amount of N and P deposited as excreta. Thus, the potential maximum amount immobilized into litter fall (needles and twigs) was estimated to account for 5-7% of total excreta-derived N and P.     

Changes in litter properties during decomposition: A study by differential thermogravimetry and scanning calorimetry

To verify the paradigm that organic matter (OM) quality (q) decreases with decomposition it is necessary to define q in strictly chemical, operational terms. We suggest defining q as the result of a balance between the energy stored in OM and the external supply of energy needed to release it. We apply this concept to the study of litter decomposition in four European pine forests: boreal, cool Atlantic, Mediterranean and warm Atlantic. Intact litter cores were taken and transported to the laboratory, where needles were sorted into six classes that summarize the main facts of the decomposition: melanisation, fragmentation and perforation by mesofauna. Each class was analyzed by both differential thermogravimetry and differential scanning calorimetry to obtain its spectra of weight loss and energy release.In the non-decomposed needles, two peaks of weight loss and energy release appear: a labile peak at about 350 °C, and a recalcitrant peak at about 450 °C. During decomposition, both peaks (but especially the recalcitrant one) move to lower temperatures, and their shapes change from well defined to flattened. In Mediterranean litters, a third peak appears at about 500 °C, due probably to refractory products of neoformation. There is a continuous increase in the energy stored in the remaining litter (in Joules per unit OM): this increase is concentrated in both the most thermolabile fractions (lost at temperatures <350 °C) and the most thermostable ones (>450 °C). With decomposition OM becomes more recalcitrant (i.e., it is lost at higher temperatures), but its stored energy becomes more available (i.e., it is released at lower temperatures). Overall, the energetic benefit/cost ratio increases. Thus, our results to date do not agree with the current paradigm that q decreases with decomposition; rather, they suggest that, at least in the first phases we studied, q is maintained or even increases.     

Degradation of hemicellulose, cellulose and lignin in decomposing spruce needle litter in relation to N

Decomposing needles from a Norway spruce forest in southern Sweden were studied for 559 days under laboratory conditions. Falling needles were collected in control (Co) plots and plots that had received 100 kg N ha⁻¹ yr⁻¹ as (NH₄)₂SO₄ for 9 years under field conditions. One of the aims was to determine whether the previously documented low decomposition rate of the N fertilized (NS) needles could be explained by a lower degradation degree of lignin. The lignin content was studied using the alkaline CuO oxidation method, the Klason lignin method and CPMAS ¹³C NMR spectroscopy. The amounts of cellulose and hemicellulose were also determined.The fertilized needle litters initially decomposed faster than the unfertilized, but later this reaction reversed, so that at the end the mass loss was 45% initial C in the control and 35% initial C in NS. Klason lignin decreased with time in both treatments and overall, the change of Klason lignin mirrored the litter mass loss. No major difference as regards the decomposition of hemicellulose occurred between the treatments, whereas significantly lower concentrations of cellulose were found in NS needles throughout the incubation. The CuO derived compounds (VSC) were somewhat lower in NS needles throughout the decomposition time. Initially, VSC increased slightly in both treatments, which contradicts the Klason lignin data. There was a weak positive relationship (p>0.05) between VSC and Klason lignin. Both vanillyls compounds (V) and cinnamyl compounds (Ci) increased slightly during decomposition, whereas syringyl compounds (S) vanished entirely. The lignin degradation degree, i.e. the acid-to-aldehyde ratio of the vanillyl compounds expressed as (Ac/Al)_v, showed no significant effect of treatment. The ¹³C NMR analyses of the combined samples showed increased content of aromatic C with increasing decomposition time. The carbohydrate content (O-alkyl C) was lower in the fertilized needle litter throughout the incubation time. The alkyl C content tended to increase with decomposition time and N fertilization. The alkyl C/O-alkyl C ratios increased in both treatments during the incubation. The NMR results were not tested statistically.In conclusion, no major difference concerning lignin degradation could be found between the unfertilized and N fertilized needle litter. Thus, the study contradicts the hypothesis that higher amounts of N reduce lignin degradation. The reduced biological activity is probably due to direct N effects on the microorganisms and their decomposing ability.     

Litter decomposition and nitrogen mineralization in newly formed gaps in a Danish beech (Fagus sylvatica) forest

Gap formation is suggested as an alternative forest management approach to avoid extreme changes in the N cycle of forest ecosystems caused by traditional management practises. The present study aimed to investigate the effect of gap formation on N availability in beech litter and mineral soil on sites, which experienced only little soil disturbance during tree harvest. N pools, litter decomposition, and N mineralization rates in mineral soil were studied in two gaps (17 and 30 m in diameter) in a 75-year-old managed European beech (Fagus sylvatica L.) forest in Denmark and related to soil temperature (5 cm depth) and soil moisture (15 cm depth). Investigations were carried out during the first 2 years after gap formation in measurement plots located along the north-south transect running through the centre of each gap and into the surrounding forest.An effect of gap size was found only for soil temperatures and litter mass loss: soil temperatures were significantly increased in the northern part of the large gap during the first year after gap formation, and litter mass loss was significantly higher in the smaller gap. All other parameters investigated revealed no effect of gap size. Nitrification, net mineralization, and soil N concentrations tended to be increased in the gaps. Cumulative rates of net mineralization were two fold higher in the gaps during the growing season (June-October), but a statistically significant increase was found only for soil NH₄-N concentrations during this period. Forest floor parameters (C:N ratios, mass loss, N release) were not significantly modified during the first year after gap formation, neither were the total C content nor the C:N ratio in mineral soil at 0-10 cm depth.     

Decomposition of beech leaves (Fagus sylvatica) and spruce needles (Picea abies) in pure and mixed stands of beech and spruce

The decomposition of spruce needles and beech leaves was investigated in a 30- and 120-yr-old beech, spruce and mixed (beech/spruce) forest using 1 mm mesh litterbags. The mass loss, content of C, N and water and microbial biomass, basal respiration and specific respiration of the litter materials were analyzed after exposure for 1.5, 3, 6, 9, 12, 18 and 24 months in the field. Decomposition of both types of litter was faster in beech than in spruce stands and after 24 months loss of C from litter materials was at a maximum in beech stands (>60%) and considerably less in the spruce and mixed stands (ca. 40%). Generally, spruce needles decomposed more rapidly than beech leaves, but the faster decay was not associated with higher N concentrations. Rather, N was accumulated more rapidly in beech leaves. Concomitantly, in beech stands microbial biomass of beech leaves exceeded that of spruce needles indicating that beech leaves consist of more favorable resources for microorganisms than spruce needles. Differences in decomposition between beech leaves and spruce needles were most pronounced in beech stands, intermediate in mixed stands and least pronounced in spruce stands. Decomposition, N content and microbial biomass in litter materials exposed in the 120-yr-old stand consistently exceeded that in the 30-yr-old stand indicating adverse conditions for litter decay in regrowing stands. Generally, mixed stands ranked intermediate between spruce and beech monocultures for most of the variables measured indicating that the adverse conditions for litter decay and microorganisms in spruce forest are effectively counteracted by admixture of beech to spruce monocultures. It is concluded that the accumulation of litter materials in spruce forests is not due to the recalcitrance of spruce needles to decay. Rather, adverse environmental conditions such as high polyphenol contents in the litter layer of spruce stands retard decomposition processes; spruce needles appear to be more sensitive to this retardation than beech leaves.     

Plant litter quality influences the contribution of soil fauna to litter decomposition in humid tropical forests, southwestern China

The aim of this field experiment was to quantify the contribution of soil fauna to plant litter decomposition in three forest sites differing in C/N ratio under natural conditions in Xishuangbanna, southwestern China. We conducted a survey of soil fauna communities, the forest floor litter and investigated mass loss of mixed tree species leaf litter for two years in a tropical secondary forest, an evergreen broad-leaf forest and a tropical rain forest. Exclusion treatments of different sized soil fauna from the leaf litter by using varying mesh size litter bags (2 mm and 0.15 mm) were also performed. Mass loss and C and N concentrations in litter bag leaf materials were determined at monthly intervals. We found that: (1) the three forests differed in floor litter biomass and nutrient contents but not in soil fauna richness and abundance; (2) litter mass loss and decomposition rate were slower when soil macrofauna and most of mesofauna were excluded; and (3) greatest soil fauna contribution to plant litter decomposition occurred in the rain forest, where leaf litter C/N ratio was also highest (41.5% contribution: 54.8 C/N ratio), in comparison to 8.69% in the broad-leaf forest and 19.52% in the secondary forest, both with low leaf litter C/N ratios (<32). Our results suggested that, soil fauna played a more pronounced role in the decomposition of mixed leaf litter in tropical rain forest, and significantly bigger effects from fauna were ascribed to the enhancement of N concentration and decrease of C concentration of the initially high C/N ratio litter in this forest site.     

Litter decomposition in primary forest and adjacent fire-disturbed forests in the Gran Sabana,southern Venezuela

Litter decomposition and nutrient dynamic were studied in tall primary forest (TF) and in adjacent slightly fire-affected (MF) and strongly fire-affected (LF) forests of the Gran Sabana, southern Venezuela. The aim of the study was to compare the mass and nutrient loss of litter in undisturbed forest and adjacent fire-disturbed forests growing under the same soil conditions. The results showed no significant differences in the dry-mass reduction among TF, MF, and LF after 1-year of decomposition. At the end of the decomposition period, the mass loss was 31% in TF, 24% in MF, and 25% in LF. With few exceptions, the initial nutrient content of the litter did not show significant differences among the studied forests. The initial litter was poor in nutrients, especially in P, with C/N ratio extremely high, particularly in LF. Both residual P content and C/N ratio were the most effective predictors of dry-mass loss. The general trend in the three studied forests was net N and to less extend P immobilization and release of K, Mg, and Ca. We concluded that during a 1-year period, the decomposition process was similar in undisturbed and adjacent fire-disturbed forests in the Gran Sabana and that the low litter decomposition seems to be mainly controlled by the low chemical quality of the decomposing litter.     

Impacts of extreme winter warming events on litter decomposition in a sub-Arctic heathland

Arctic climate change is expected to lead to a greater frequency of extreme winter warming events. During these events, temperatures rapidly increase to well above 0 °C for a number of days, which can lead to snow melt at the landscape scale, loss of insulating snow cover and warming of soils. However, upon return of cold ambient temperatures, soils can freeze deeper and may experience more freeze-thaw cycles due to the absence of a buffering snow layer. Such loss of snow cover and changes in soil temperatures may be critical for litter decomposition since a stable soil microclimate during winter (facilitated by snow cover) allows activity of soil organisms. Indeed, a substantial part of fresh litter decomposition may occur in winter. However, the impacts of extreme winter warming events on soil processes such as decomposition have never before been investigated. With this study we quantify the impacts of winter warming events on fresh litter decomposition using field simulations and lab studies.Winter warming events were simulated in sub-Arctic heathland using infrared heating lamps and soil warming cables during March (typically the period of maximum snow depth) in three consecutive years of 2007, 2008, and 2009. During the winters of 2008 and 2009, simulations were also run in January (typically a period of shallow snow cover) on separate plots. The lab study included soil cores with and without fresh litter subjected to winter-warming simulations in climate chambers.Litter decomposition of common plant species was unaffected by winter warming events simulated either in the lab (litter of Betula pubescens ssp. czerepanovii), or field (litter of Vaccinium vitis-idaea, and B. pubescens ssp. czerepanovii) with the exception of Vaccinium myrtillus (a common deciduous dwarf shrub) that showed less mass loss in response to winter warming events. Soil CO₂ efflux measured in the lab study was (as expected) highly responsive to winter warming events but surprisingly fresh litter decomposition was not. Most fresh litter mass loss in the lab occurred during the first 3-4 weeks (simulating the period after litter fall).In contrast to past understanding, this suggests that winter decomposition of fresh litter is almost non-existent and observations of substantial mass loss across the cold season seen here and in other studies may result from leaching in autumn, prior to the onset of “true” winter. Further, our findings surprisingly suggest that extreme winter warming events do not affect fresh litter decomposition.     

Nutrient limitation of litter decomposition with long-term secondary succession: evidence from controlled laboratory experiments

Purpose

Understanding ecosystem processes such as litter decomposition in response to dramatic land-use change is critical for modeling and predicting carbon (C) cycles. However, the patterns of litter decomposition along with long-term secondary succession (over 100 years) are not well reported, especially concerning nutrient limitations on litter decomposition.

Materials and methods

To clarify the response of litter decomposition to changes in soil nutrient availability, we conducted four incubation experiments involving soil and litter and nutrient addition from different successional stages and investigated the changes in microbial respiration and litter mass loss.

Results and discussion

Our results revealed that microbial respiration increased with succession without any litter addition (1.19~1.73 mg C g^?1 soil), and litter addition significantly promoted microbial respiration (16.5~72.9%), especially in the early successional stage (grassland and shrubland). The decomposition rate of the same litter decreased with succession. In addition, nitrogen (N) and phosphorus (P) addition showed significant effects on litter decomposition and microbial respiration; P addition promoted litter decomposition (2.4~15.3%) and microbial respiration (10.1~34.5%) in all successional stages, while N addition promoted litter decomposition (4.0~10.3%) and microbial respiration (5.4~27.2%) in all except the last stage of succession, which showed a negative effect on litter decomposition (??7.5%) and microbial respiration (??6.1%), indicating possible N saturation of litter decomposition and microbial respiration.

Conclusions

This work highlights that soil nutrient availability and successional stages need to be taken into account to predict the changes to litter decomposition in response to global changes.

     

The effect of temperature on decomposition of leaf litter from two tropical forests by a microcosm experiment

A 120 days’ incubation experiment was conducted to analyze the effect of temperature on the decomposition of leaf litter (Altingia obovata) in two tropical primary montane rainforests with different precipitation conditions. The results showed no difference in mass loss of leaf litter between the two forests at 20 °C, in spite that Jianfengling forest had less precipitation than Diaoluoshan forest. But higher mass loss of leaf litter was found from Jianfengling forest site (30.1%) than that from Diaoluoshan forest site (25.9%) at 30 °C at the end of incubation. Lignin exhibited higher mass loss from Jianfengling forest (29.9%) than from Diaoluoshan forest (23.3%) at 20 °C, but no difference between two forest sites at 30 °C. Total carbohydrates were decomposed faster by the decomposers from Diaoluoshan forest site (42.7%) than that from Jianfengling forest site (36.3%) at 20 °C, but 46.6% and 38.5% for Jianfengling and Diaoluoshan montane rainforests, respectively, at 30 °C. Temperature increase did not significantly lead to the difference in mass loss of leaf litter for the two forest sites. Temperature increase did not affect lignin loss for Diaoluoshan forest, but reduced lignin loss for Jianfengling forest. Temperature increase accelerated the decomposition of carbohydrate for Jianfengling forest, but opposite for Diaoluoshan forest. The response of decomposition of leaf litter to forest type and temperature was positively related to the difference in microbial activities between both montane rainforests.     

A litter-slurry technique elucidates the key role of enzyme production and microbial dynamics in temperature sensitivity of organic matter decomposition

The rate at which organic matter decomposes generally increases with temperature, unless it is physico-chemically protected from enzymatic depolymerization. The temperature sensitivity of decomposition should increase with decreasing reaction rates, corresponding to increasing activation energy of the decomposing compounds. One approach to testing this carbon-quality temperature hypothesis is to study the effect of temperature on leaf litter decomposition, because fresh surface litter is unprotected. However, other factors such as humidity co-vary with temperature, and biological processes such as enzyme production and microbial population growth may also be thermally sensitive. We developed a litter slurry approach to isolate the effect of temperature and litter quality on decomposition. We found that pine litter decomposed faster than oak litter, consistent with a lower C:N and lignin:N ratio. During the first 14 days of decomposition, there was no difference in decomposition rate for litter incubated at 25 °C compared to 35 °C. Lower potential enzyme activity at 35 °C suggested that enzyme production was suppressed at 35 °C compared to 25 °C, resulting in similar in situ enzyme activities at the two temperatures. After 14 days, enzyme pools were similar between the two incubation temperatures, which resulted in faster decomposition at the warmer temperature, consistent with enzyme kinetic theory. At Day 14, the decomposition rate of the high quality pine litter was more temperature sensitive than the decomposition rate of the lower quality oak litter, suggesting that the quality of soluble pool rather than bulk chemistry determined the temperature sensitivity during this stage. After 28 days of incubation, oak litter decomposition was more temperature sensitive than pine litter, consistent with the carbon temperature-quality hypothesis. The litter slurry approach revealed that biological responses to temperature can affect the apparent temperature sensitivity of decomposition, and highlight a need for further research into microbial responses to temperature.     

Home-field advantage of litter decomposition and nitrogen release in forest ecosystems

Litter decomposition is a major fundamental ecological process that regulates nutrient cycling, thereby affecting net ecosystem carbon (C) storage as well as primary productivity in forest ecosystems. Litter decomposes in its home environment faster than in any other environment. However, evidence for this phenomenon, which is called the home-field advantage (HFA), has not been universal. We provide the first HFA quantification of litter decomposition and nutrient release through meta-analysis of published data in global forest ecosystems. Litter mass loss was 4.2 % faster on average, whereas nitrogen (N) release was 1.7 % lower at the home environment than in another environment. However, no HFA of phosphorus (P) release was observed. Broadleaf litter (4.4 %) had a higher litter mass loss HFA than coniferous litter (1.0 %). The positive HFA of N release was found in the coniferous litter. Mass loss HFA was significantly and negatively correlated with the initial lignin:N litter ratio. The litter decomposition and N release HFAs were obtained when mesh size ranged from 0.15 mm to 2.0 mm. The HFA of litter decomposition increased with decomposition duration during the early decomposition stage. The HFA of N release was well correlated with mass loss, and the greatest HFA was at mass loss less than 20 %. Our results suggest that the litter decomposition and N release HFAs are widespread in forest ecosystems. Furthermore, soil mesofauna is significantly involved in the HFA of litter decomposition.     

Enzyme activities and litter decomposition in agricultural soils in northern,central, and southern Germany

Although enzyme activities were extensively investigated in soils with reference to abiotic environmental conditions and human impact, their role in litter decomposition is not fully understood. Therefore, decomposition rates and enzyme activities were studied using nylon bags and three litter types buried in silty‐loamy Cambisols and Luvisols in northern, central, and southern Germany under similar averaged temperature and precipitation and a maritime to continental gradient. After 180 d, the ash‐free mass remaining ranged between 15% and 68% for the Triticum, Secale, and Lolium litter. The enzyme activities were mainly controlled by the litter type and the decomposition time and less but significant by site. The highest decomposition rate occurred at the central German site for Lolium litter associated with highest arginine ammonification and urease activity in litter. In contrast, the recalcitrant Secale and Triticum litter were decomposed more rapidly at the northern and southern site where urease, protease, and arginine ammonification was high in the bulk soil. The β‐glucosidase activity was similar in soil and litter at the three locations and was not correlated to the velocity of litter decomposition. Since the abiotic environmental factors at the maritime to continental gradient did not explain the site‐specific velocity of both rapidly decomposing and refractory litter, enzyme activities related to the N cycling like arginine ammonification and urease activity were recognized to velocity of litter decomposition.     

Photodegradation of plant litter in the Sonoran Desert varies by litter type and age

Predicting litter decay rates in arid systems has proved elusive and sunlight (photodegradation) is a potentially important but poorly understood driver of litter decay in these systems. We placed three litter types (Cynodon dactylon, Larrea tridentata leaves, and L. tridentata twigs) in envelopes whose tops either transmitted all solar radiation, filtered UV-B, filtered all UV, or filtered all UV and visible solar radiation, on the soil surface of the Sonoran Desert and assessed mass loss over 14 months. Regardless of treatment, final mass loss was greatest in C. dactylon litter and least in L. tridentata twig litter, consistent with initial litter characteristics of presumed litter quality; C. dactylon had the lowest lignin concentration and lignin:N, and the highest cellulose:lignin and area:mass. Compared to litter in sunlight, excluding solar UV, or UV-B, slowed mass loss of all 3 litter types, and UV-B appeared more effective than UV-A in photodegradation. The relative contribution of UV photodegradation to mass loss increased with litter age. After 14 months, litter exposed to solar UV lost 1.2 (C. dactylon), 1.3 (L. tridentata twigs) and 1.4 (L. tridentata leaves) times as much mass as litter not exposed to UV radiation. The relative contribution of UV photodegradation to mass loss increased with the initial C:N ratio of litter, but was not related to initial lignin concentration or optical properties (i.e. UV and visible absorbance and transmittance) of litter. Within all litter type by treatment combinations, there was a strong positive correlation between litter mass loss and ash concentration. In some cases, a discontinuity in this relationship was detected, suggesting a threshold ash concentration, above which further mass loss was negligible. We expected these thresholds to be most prevalent in sunlight, because soil films could prevent sunlight from reaching litter and thereby minimize photodegradation. Contrary to expectations, thresholds were more common in shade or UV filter treatments, suggesting that reductions in photodegradation attributable to soil films were not typically responsible. The effect of shading, which likely enhanced microbial degradation via higher relative humidity due to lower temperatures, depended on litter type and time. Compared to litter in sunlight, mass loss of shaded litter was greater over the initial 3 months in all litter types, illustrating that microbial degradation in shade was greater than photodegradation in sunlight. These differences in mass loss between shaded and sunlit litter increased over the 14 month experiment in L. tridentata twigs, declined in L. tridentata leaves, and disappeared within 6 months in C. dactylon, illustrating that the timing of this shift in the dominance of photodegradation versus microbial degradation was highly dependent on litter type. In a second experiment, we reduced microclimate differences among sun and shade treatments, pre-sterilized litter to reduce microbial degradation, and examined the mass loss of young and old and L. tridentata leaf litter after 53 days outdoors. Consistent with our first experiment, mass loss attributable to photodegradation was greater in old than young litter. Unsterilized litter exposed to sunlight (UV and visible) lost 1.3 (young) and 1.5 (old) times as much mass as shaded litter. Pre-sterilized litter exposed to sunlight lost 11.4 (young litter) and 45.9 (old litter) times as much mass as shaded litter. These large differences in pre-sterilized litter were the result of the very small mass loss of shaded litter (≤0.2%), together with modest losses of sunlit litter (<5%). Taken together, our findings suggest that as litter aged, microbial degradation became a weaker driver of mass loss, while photodegradation became stronger.     

Multiple effects of reindeer grazing on the soil processes in nutrient-poor northern boreal forests

Reindeer grazing has a great influence on the ground vegetation of nutrient-poor northern boreal forests dominated by Cladonia lichens in Fennoscandia. Grazing may influence the soil processes in these systems either by influencing the quality of plant litter, or by indirect effects through the soil microclimate. In order to investigate the mechanisms underlying the effects of reindeer on boreal forest soils, we analyzed litter decomposition, soil and microbial C and N, microbial community composition, and soil organic matter quality in three forest sites with old reindeer exclosures adjacent to grazed areas. There was no effect of grazing on soil C/N ratio, inorganic N concentrations, microbial biomass C, microbial community structure analyzed by phospholipid fatty acid (PLFA) analysis, and organic matter quality analyzed by sequential fractionation, in the soil organic layer. However, microbial N was enhanced by grazing at some of the sampling dates and was negatively correlated with soil moisture, which indicates that increased microbial N could be a stress response to drought. The effect of grazing on litter decomposition varied among the decomposition stages: during the first 1.5 months, the litter C loss was significantly higher in the grazed than the ungrazed areas, but the difference rapidly levelled out and, after one year, the accumulated litter C loss was higher in the ungrazed than the grazed areas. Litter N loss was, however, higher in the grazed areas. Our study demonstrates that herbivores may influence soil processes through several mechanisms at the same time, and to a varying extent in the different stages of decomposition.     

Scots pine litter decomposition along drainage succession and soil nutrient gradients in peatland forests, and the effects of inter-annual weather variation

Peatlands form a large carbon (C) pool but their C sink is labile and susceptible to changes in climate and land-use. Some pristine peatlands are forested, and others have the potential: the amount of arboreal vegetation is likely to increase if soil water levels are lowered as a consequence of climate change. On those sites tree litter dynamics may be crucial for the C balance. We studied the decomposition of Scots pine (Pinus sylvestris L.) needle and root litter in boreal peatland sites representing gradients in drainage succession (succession following water level drawdown caused by forest drainage) and soil nutrient level during several years of varying weather conditions. Neither gradient had an unambiguous effect on litter mass loss. Mass loss over 2 years was faster in undrained versus drained sites for both needle litter, incubated in the moss layer, and fine root litter, incubated in 0-10 cm peat layer, suggesting moisture stress in the surface layers of the drained sites limited decomposition. Differences among the drained sites were not consistent. Among years, mass loss correlated positively with precipitation variables, and mostly negatively or not at all with temperature sum. We concluded that a long-term water level drawdown in peatlands does not necessarily enhance decay of fresh organic matter. Instead, the drained site may turn into a ‘large hummock-system’ where several factors, including litter quality, relative moisture deficiency, higher acidity, lower substrate temperature, and in deeper layers also oxygen deficiency, may interact to constrain organic matter decomposition. Further, the decomposition rates may not vary systematically among sites of different soil nutrient levels following water level drawdown. Our results emphasize the importance of annual weather variations on decomposition rates, and demonstrate that single-period incubation studies incorporate an indeterminable amount of temporal variation.     

Carbon and nitrogen turnover in response to warming and nitrogen addition during early stages of forest litter decomposition—an incubation experiment

Purpose

Little is known about the interactive effects of temperature, nitrogen (N) supply, litter quality, and decomposition time on the turnover of carbon (C) and N of forest litter. The objective of this study was to investigate the interactive effects of warming, N addition and tree species on the turnover of C and N during the early decomposition stage of litters in a temperate forest.

Materials and methods

A 12-week laboratory incubation experiment was carried out. The leaf litters including two types of broadleaf litters (Quercus mongolica and Tilia amurensis), a needle litter (Pinus koraiensis), and a mixed litter of them were collected from a broad-leaved Korean pine mixed forest ecosystem in northeastern China in September 2009. Nine treatments were conducted using three temperatures (15, 25, and 35 °C) combined with three doses of N addition (equal to 0, 75, and 150 kg?·?ha^?1?a^?1, respectively, as NH₄NO₃).

Results and discussion

After 12 weeks of incubation, the mass loss ranged between 12 and 35 %. The broadleaf litters had greater mass loss and cumulative CO₂–C emission than the needle litter. Temperature and N availability interacted to affect litter mass loss and decomposition rate. The dissolved organic carbon (DOC) and nitrogen (DON) concentrations in litter leachate varied widely with litter types. DOC increased significantly with increased temperature but decreased significantly with increased N availability. DON increased significantly with increased N availability but showed a higher level at the moderate decomposition temperature. The amounts of CO₂ and N₂O emission were significantly higher at 25 °C than those at 15 and 35 °C, and were significantly increased by the N addition.

Conclusions

The present study indicated relatively intricate temperature and N addition effects on C and N cycling during early stages of litter decomposition, implying that future increases in temperature and N deposition will directly affect C and N cycling in broad-leaved Korean pine mixed forest ecosystem, and may indirectly influence the ecosystem composition, productivity, and functioning in NE China. It is, therefore, important to understand the interactive effects of biotic and abiotic factors on litter decomposition in field conditions in order to assess and predict future ecosystem responses to environmental changes in NE China.     

Effects of surface fire on litter decomposition and occurrence of microfungi in a Cymbopogon polyneuros dominated grassland

Abstract

Litter decomposition rate, changes in macronutrients such as nitrogen (N), phosphorus (P) and potassium (K) from different grades of litter decomposition and occurrence of soil microfungi were investigated in a Cymbopogon polyneuros-dominated tall grass ecosystem from a burned and an unburned site in southern India. The litter decomposition rate was higher at the burned site than at the unburned site. This rate was also higher when the litter was mixed with the mineral soil material than leaving the litter unaffected on the soil surface. The concentrations of N, P, and K in the litter decreased as a result of progressed litter decomposition. Occurrence of microfungi identified from the different decomposition grades of the Cymbopogon polyneuros litter was higher at the burned site compared to the unburned site. Microfungal species present at both sites showed only minor differences.     

Effects of nitrogen addition and litter properties on litter decomposition and enzyme activities of individual fungi

Litter decomposition is an important process of C and N cycling in the soil. Variation in the response of litter decomposition to nitrogen (N) addition (positive, negative or neutral) has been observed in many field studies. However, mechanism about variability in individual fungal species response to N addition has not yet been well demonstrated in the literature. Therefore, the objective of this study was to investigate the effects of N addition and litter chemistry properties on litter decomposition and enzyme activities of individual fungi. Three fungal species (Penicillium, Aspergillus, and Trichoderma) were isolated from a subtropical mixed forest soil. An incubation experiment was conducted using the individual fungi with two types of litter (leaf of Pinus massoniana and needle of Cryptocarya chinensis) and different N addition levels (0, 50 and 100 for N-deficient treatments, and 500 and 1000 μg N for N-excessive treatments). Cumulative CO₂-C, enzyme activities, and lignin and cellulose loss were measured during the incubation period of 60 days. Litter decomposition and enzyme activities significantly varied with the fungal species, while the N addition and litter types greatly affected fungal enzyme activities. The N treatments significantly increased lignin-rich needle decomposition by lignocellulose decomposers (Penicillium and Aspergillus) but did not affect their leaf decomposition. On the contrary, The N treatments stimulated leaf decomposition by cellulolytic species (Trichoderma) but did not affect its needle decomposition. Correlation analysis showed that lignin in the litter was the key component to affect litter decomposition. Activities of N-acetyl-β-glucosaminidase and phenol oxidase were both positively correlated to litter decomposition. The fungi (Penicillium and Aspergillus) with higher production of N-acetyl-β-glucosaminidase showed higher litter decomposition ability. The low N addition levels stimulated Penicillium and Aspergillus litter decomposition, but they still required more N source (e.g., litter N source) to support decomposition. Depressed fungal litter N uptake (lower N-acetyl-β-glucosaminidase activities) only occurred at the highest N addition level. Litter decomposition of Trichoderma depended more on external N and its litter decomposition capability was the lowest among the three species.