Abstract. We maintained a factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment for 11 years in a humid lowland forest growing on a relatively fertile soil in Panama to evaluate potential nutrient limitation of tree growth rates, fine-litter production, and fine-root biomass. We replicated the eight factorial treatments four times using 32 plots of 40 3 40 m each. The addition of K was associated with significant decreases in stand-level fineroot biomass and, in a companion study of seedlings, decreases in allocation to roots and increases in height growth rates. The addition of K and N together was associated with significant increases in growth rates of saplings and poles (1-10 cm in diameter at breast height) and a further marginally significant decrease in stand-level fine-root biomass. The addition of P was associated with a marginally significant (P ¼ 0.058) increase in fine-litter production that was consistent across all litter fractions. Our experiment provides evidence that N, P, and K all limit forest plants growing on a relatively fertile soil in the lowland tropics, with the strongest evidence for limitation by K among seedlings, saplings, and poles.

The widespread use of forest litter as animal bedding in central Europe for many centuries gave rise to the first litter manipulation studies, and their results demonstrated that litter and its decomposition are a vital part of ecosystem function. Litter plays two major roles in forest ecosystems: firstly, litterfall is an inherent part of nutrient and carbon cycling, and secondly, litter forms a protective layer on the soil surface that also regulates microclimatic conditions. By reviewing 152 years of litter manipulation experiments, I show that the effects of manipulating litter stem from changes in one, or both, of these two functions, and interactions between the variables influenced by the accumulation of litter can result in feedback mechanisms that may intensify treatment effects or mask responses, making the interpretation of results difficult.Long-term litter removal increased soil bulk density, overland flow, erosion, and temperature fluctuations and upset the soil water balance, causing lower soil water content during dry periods. Soil pH increased or decreased in response to manipulation treatments depending on forest type and initial soil pH, but it is unclear why there was no uniform response. Long-term litter harvesting severely depleted the forests of nutrients. Decreases in the concentrations of available P, Ca, Mg, and K in the soil occurred after only three to five years. The decline in soil N occurred over longer periods of time, and the relative loss was greater in soils with high initial nitrogen concentration. Tree growth declined with long-term litter removal, probably due to lower nutrient availability. Litter manipulation also added or removed large amounts of carbon thereby affecting microbial communities and altering soil respiration rates.Litter manipulation experiments have shown that litter cover acts as a physical barrier to the shoot emergence of small-seeded species ; further, the microclimate maintained by the litter layer may be favourable to herbivores and pathogens and is important in determining later seedling survival and performance. Litter manipulation altered the competitive outcomes between tree seedlings and forbs, thereby influencing species composition and diversity ; changes in the species composition of understorey vegetation following treatments occurred fairly rapidly. By decreasing substrate availability and altering the microclimate, litter removal changed fungal species composition and diversity and led to a decline in populations of soil fauna. However, litter addition did not provoke a corresponding increase in the abundance or diversity of fungi or soil fauna.Large-scale long-term studies are still needed in order to investigate the interactions between the many variables affected by litter, especially in tropical and boreal forests, which have received little attention. Litter manipulation treatments present an opportunity to assess the effects of increasing primary production in forest ecosystems; specific research aims include assessing the eff...

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Aboveground litter production in forests is likely to increase as a consequence of elevated atmospheric carbon dioxide (CO2) concentrations, rising temperatures, and shifting rainfall patterns. As litterfall represents a major flux of carbon from vegetation to soil, changes in litter inputs are likely to have wide-reaching consequences for soil carbon dynamics. Such disturbances to the carbon balance may be particularly important in the tropics because tropical forests store almost 30% of the global soil carbon, making them a critical component of the global carbon cycle; nevertheless, the effects of increasing aboveground litter production on belowground carbon dynamics are poorly understood. We used long-term, large-scale monthly litter removal and addition treatments in a lowland tropical forest to assess the consequences of increased litterfall on belowground CO2 production. Over the second to the fifth year of treatments, litter addition increased soil respiration more than litter removal decreased it; soil respiration was on average 20% lower in the litter removal and 43% higher in the litter addition treatment compared to the controls but litter addition did not change microbial biomass. We predicted a 9% increase in soil respiration in the litter addition plots, based on the 20% decrease in the litter removal plots and an 11% reduction due to lower fine root biomass in the litter addition plots. The 43% measured increase in soil respiration was therefore 34% higher than predicted and it is possible that this ‘extra’ CO2 was a result of priming effects, i.e. stimulation of the decomposition of older soil organic matter by the addition of fresh organic matter. Our results show that increases in aboveground litter production as a result of global change have the potential to cause considerable losses of soil carbon to the atmosphere in tropical forests.

Summary1. The cycling of nutrients in litterfall is considered a key mechanism in the maintenance of tropical forest fertility but its importance has rarely been quantified experimentally. 2. We carried out a long-term (5 years), large-scale litter manipulation experiment in lowland semievergreen tropical forest to determine how changes in litterfall affect forest nutrient cycling. We hypothesized that: (i) long-term litter removal would decrease the forest's nutrient supply; (ii) litter addition would increase the forest's nutrient supply; (iii) soil and foliar nutrient concentrations would change in response to litter manipulation and would eventually affect above-ground productivity. 3. To test our hypotheses, we measured trunk growth, litterfall, and nutrient concentrations in live leaves, litter and soil in plots where litter was removed once a month (L)), litter was added once a month (L+) and controls (CT). 4. After 5 years, the concentration of nitrate in the soil and soil stocks of inorganic nitrogen were higher in the L+plots and lower in the L) plots compared to the controls. Ammonium concentrations in the soil were also lower in the L) plots. Nitrogen in leaves and litter and the annual nitrogen return by litter were higher in the L+plots, while potassium return was lower in the L) plots. Surprisingly, our treatments had little effect on phosphorus in soil, leaves or litter, even though lowland tropical forests are generally thought to be largely phosphorus limited. 5. Trunk growth of large trees was not affected by litter manipulation but rainy season litterfall from 2003 to 2008 was 13% higher in the L+plots compared to the controls. 6. Synthesis. Litter removal affected forest nutrient cycling and productivity less than expected, probably because the soil at our site is moderately fertile. However, litter addition increased litterfall indicating that some limitation of forest production was removed by litter addition. We expected strong effects of litter manipulation on phosphorus cycling; instead, we found a stronger effect on nitrogen cycling. Our results suggest that litter is an important source of nutrients, in particular nitrogen, to trees in this lowland semi-evergreen tropical forest.

Abstract. Global change has been shown to alter the amount of above-ground litter inputs to soil greatly, which could cause substantial cascading effects on below-ground biogeochemical cycling. Despite extensive study, there is uncertainty about how changes in above-ground litter inputs affect soil carbon and nutrient turnover and transformation. Here, we conducted a meta-analysis on 70 litter-manipulation experiments in order to assess how changes in above-ground litter inputs alter soil physicochemical properties, carbon dynamics and nutrient cycles. Our results demonstrated that litter removal decreased soil respiration by 34 %, microbial biomass carbon in the mineral soil by 39 % and total carbon in the mineral soil by 10 %, whereas litter addition increased them by 31, 26 and 10 %, respectively. This suggests that greater litter inputs increase the soil carbon sink despite higher rates of carbon release and transformation. Total nitrogen and extractable inorganic nitrogen in the mineral soil decreased by 17 and 30 %, respectively, under litter removal, but were not altered by litter addition. Overall, litter manipulation had a significant impact upon soil temperature and moisture, but not soil pH; litter inputs were more crucial in buffering soil temperature and moisture fluctuations in grassland than in forest. Compared to other ecosystems, tropical and subtropical forests were more sensitive to variation in litter inputs, as altered litter inputs affected the turnover and accumulation of soil carbon and nutrients more substantially over a shorter time period. Our study demonstrates that although the magnitude of responses differed greatly among ecosystems, the direction of the responses was very similar across different ecosystems. Interactions between plant productivity and below-ground biogeochemical cycling need to be taken into account to predict ecosystem responses to environmental change.

Root proliferation into the Oa and Oe soil horizons in tropical forests is often substantial and allows direct cycling of nutrients from the organic matter; this was thought to be an adaptation to the low nutrient supply in infertile soils. In this study, we show that experimentally increased litter inputs promote root proliferation into the Oi and Oe horizons in a relatively fertile soil, suggesting that it is a response to a more readily available nutrient source rather than an adaptation to nutrient shortage, and the absence of root mats on fertile tropical soils is simply a consequence of the lack of persistent organic horizons due to high decomposition rates.

Predicting future impacts of anthropogenic change on tropical forests requires a clear understanding of nutrient constraints on productivity. We compared experimental fertilization and litter manipulation treatments in an old-growth lowland tropical forest to distinguish between the effects of inorganic nutrient amendments and changes in nutrient cycling via litterfall. We measured the changes in soil and litter nutrient pools, litterfall, and fine root biomass in plots fertilized with nitrogen (N), phosphorus (P), or potassium (K), and in litter addition and litter removal treatments during 7 years. Soil inorganic N and litter N increased in double-litter plots but not in N-fertilized plots. Conversely, fitter P and soil pools of P and K increased in fertilized plots but not in the double-litter plots. Soil and litter pools of N and K decreased in the no-litter plots.Changes in litterfall with added nutrients or litter were only marginally significant, but fine root biomass decreased with both the litter and the K addition. Differences between the two experiments are mostly attributable to the coupled cycling of carbon and nutrients in litter. Increased nutrient inputs in litter may improve plant uptake of some nutrients compared to fertilization with similar amounts. The litter layer also appears to play a key role in nutrient retention. We discuss our findings in the context of possible impacts of anthropogenic change on tropical forests.