Arbuscular mycorrhizal fungal community composition is altered by long-term litter removal but not litter addition in a lowland tropical forest

Tropical forest productivity is sustained by the cycling of nutrients through decomposing organic matter. Arbuscular mycorrhizal (AM) fungi play a key role in the nutrition of tropical trees, yet there has been little experimental investigation into the role of AM fungi in nutrient cycling via decomposing organic material in tropical forests. We evaluated the responses of AM fungi in a long-term leaf litter addition and removal experiment in a tropical forest in Panama. We described AM fungal communities using 454-pyrosequencing, quantified the proportion of root length colonised by AM fungi using microscopy, and estimated AM fungal biomass using a lipid biomarker. AM fungal community composition was altered by litter removal but not litter addition. Root colonisation was substantially greater in the superficial organic layer compared with the mineral soil. Overall colonisation was lower in the litter removal treatment, which lacked an organic layer. There was no effect of litter manipulation on the concentration of the AM fungal lipid biomarker in the mineral soil. We hypothesise that reductions in organic matter brought about by litter removal may lead to AM fungi obtaining nutrients from recalcitrant organic or mineral sources in the soil, besides increasing fungal competition for progressively limited resources

Arbuscular mycorrhizal fungal community composition is altered by long-term litter removal but not litter addition in a lowland tropical forest

Tropical forest productivity is sustained by the cycling of nutrients through decomposing organic matter. Arbuscular mycorrhizal (AM) fungi play a key role in the nutrition of tropical trees, yet there has been little experimental investigation into the role of AM fungi in nutrient cycling via decomposing organic material in tropical forests. We evaluated the responses of AM fungi in a long-term leaf litter addition and removal experiment in a tropical forest in Panama. We described AM fungal communities using 454-pyrosequencing, quantified the proportion of root length colonised by AM fungi using microscopy, and estimated AM fungal biomass using a lipid biomarker. AM fungal community composition was altered by litter removal but not litter addition. Root colonisation was substantially greater in the superficial organic layer compared with the mineral soil. Overall colonisation was lower in the litter removal treatment, which lacked an organic layer. There was no effect of litter manipulation on the concentration of the AM fungal lipid biomarker in the mineral soil. We hypothesise that reductions in organic matter brought about by litter removal may lead to AM fungi obtaining nutrients from recalcitrant organic or mineral sources in the soil, besides increasing fungal competition for progressively limited resources.Smithsonian Tropical Research Institute; Cambridge Home and European Scholarship; Department of Plant Sciences, Cambridge; Cambridge Philosophical Society; European Research Council; European Union's Seventh Framework Programme. Grant Number: FP/2007-201

Plant responses to fertilization experiments in lowland, species-rich, tropical forests.

We present a meta-analysis of plant responses to fertilization experiments conducted in lowland, species-rich, tropical forests. We also update a key result and present the first species-level analyses of tree growth rates for a 15-yr factorial nitrogen (N), phosphorus (P), and potassium (K) experiment conducted in central Panama. The update concerns community-level tree growth rates, which responded significantly to the addition of N and K together after 10 yr of fertilization but not after 15 yr. Our experimental soils are infertile for the region, and species whose regional distributions are strongly associated with low soil P availability dominate the local tree flora. Under these circumstances, we expect muted responses to fertilization, and we predicted species associated with low-P soils would respond most slowly. The data did not support this prediction, species-level tree growth responses to P addition were unrelated to species-level soil P associations. The meta-analysis demonstrated that nutrient limitation is widespread in lowland tropical forests and evaluated two directional hypotheses concerning plant responses to N addition and to P addition. The meta-analysis supported the hypothesis that tree (or biomass) growth rate responses to fertilization are weaker in old growth forests and stronger in secondary forests, where rapid biomass accumulation provides a nutrient sink. The meta-analysis found no support for the long-standing hypothesis that plant responses are stronger for P addition and weaker for N addition. We do not advocate discarding the latter hypothesis. There are only 14 fertilization experiments from lowland, species-rich, tropical forests, 13 of the 14 experiments added nutrients for five or fewer years, and responses vary widely among experiments. Potential fertilization responses should be muted when the species present are well adapted to nutrient-poor soils, as is the case in our experiment, and when pest pressure increases with fertilization, as it does in our experiment. The statistical power and especially the duration of fertilization experiments conducted in old growth, tropical forests might be insufficient to detect the slow, modest growth responses that are to be expected

Responses of arbuscular mycorrhizal fungi to long-term inorganic and organic nutrient addition in a lowland tropical forest

Improved understanding of the nutritional ecology of arbuscular mycorrhizal (AM) fungi is important in understanding how tropical forests maintain high productivity on low-fertility soils. Relatively little is known about how AM fungi will respond to changes in nutrient inputs in tropical forests, which hampers our ability to assess how forest productivity will be influenced by anthropogenic change. Here we assessed the influence of long-term inorganic and organic nutrient additions and nutrient depletion on AM fungi, using two adjacent experiments in a lowland tropical forest in Panama. We characterised AM fungal communities in soil and roots using 454-pyrosequencing, and quantified AM fungal abundance using microscopy and a lipid biomarker. Phosphorus and nitrogen addition reduced the abundance of AM fungi to a similar extent, but affected community composition in different ways. Nutrient depletion (removal of leaf litter) had a pronounced effect on AM fungal community composition, affecting nearly as many OTUs as phosphorus addition. The addition of nutrients in organic form (leaf litter) had little effect on any AM fungal parameter. Soil AM fungal communities responded more strongly to changes in nutrient availability than communities in roots. This suggests that the 'dual niches' of AM fungi in soil versus roots are structured to different degrees by abiotic environmental filters, and biotic filters imposed by the plant host. Our findings indicate that AM fungal communities are fine-tuned to nutrient regimes, and support future studies aiming to link AM fungal community dynamics with ecosystem function

Changes in soil carbon and nutrients following six years of litter removal and addition in a tropical semi-evergreen rain forest

Abstract. Increasing atmospheric CO2 and temperature may increase forest productivity, including litterfall, but the consequences for soil organic matter remain poorly understood. To address this, we measured soil carbon and nutrient concentrations at nine depths to 2 m after six years of continuous litter removal and litter addition in a semi-evergreen rain forest in Panama. Soils in litter addition plots, compared to litter removal plots, had higher pH and contained greater concentrations of KCl-extractable nitrate (both to 30 cm); Mehlich-III extractable phosphorus and total carbon (both to 20 cm); total nitrogen (to 15 cm); Mehlich-III calcium (to 10 cm); Mehlich-III magnesium and lower bulk density (both to 5 cm). In contrast, litter manipulation did not affect ammonium, manganese, potassium or zinc, and soils deeper than 30 cm did not differ for any nutrient. Comparison with previous analyses in the experiment indicates that overall the effect of litter manipulation on nutrient concentrations and the depth to which the effects are significant are increasing with time. To allow for changes in bulk density in calculation of changes in carbon stocks, we standardized total carbon and nitrogen on the basis of a constant mineral mass. For 200 kg m−2 of mineral soil (approximately the upper 20 cm of the profile) about 0.5 kg C m−2 was 'missing' from the litter removal plots, with a similar amount accumulated in the litter addition plots. There was an additional 0.4 kg C m−2 extra in the litter standing crop of the litter addition plots compared to the control. This increase in carbon in surface soil and the litter standing crop can be interpreted as a potential partial mitigation of the effects of increasing CO2 concentrations in the atmosphere. </jats:p

Changes in soil carbon and nutrients following 6 years of litter removal and addition in a tropical semi-evergreen rain forest

Abstract. Increasing atmospheric CO2 and temperature may increase forest productivity, including litterfall, but the consequences for soil organic matter remain poorly understood. To address this, we measured soil carbon and nutrient concentrations at nine depths to 2 m after 6 years of continuous litter removal and litter addition in a semi-evergreen rain forest in Panama. Soils in litter addition plots, compared to litter removal plots, had higher pH and contained greater concentrations of KCl-extractable nitrate (both to 30 cm); Mehlich-III extractable phosphorus and total carbon (both to 20 cm); total nitrogen (to 15 cm); Mehlich-III calcium (to 10 cm); and Mehlich-III magnesium and lower bulk density (both to 5 cm). In contrast, litter manipulation did not affect ammonium, manganese, potassium or zinc, and soils deeper than 30 cm did not differ for any nutrient. Comparison with previous analyses in the experiment indicates that the effect of litter manipulation on nutrient concentrations and the depth to which the effects are significant are increasing with time. To allow for changes in bulk density in calculation of changes in carbon stocks, we standardized total carbon and nitrogen on the basis of a constant mineral mass. For 200 kg m−2 of mineral soil (approximately the upper 20 cm of the profile) about 0.5 kg C m−2 was “missing” from the litter removal plots, with a similar amount accumulated in the litter addition plots. There was an additional 0.4 kg C m−2 extra in the litter standing crop of the litter addition plots compared to the control. This increase in carbon in surface soil and the litter standing crop can be interpreted as a potential partial mitigation of the effects of increasing CO2 concentrations in the atmosphere.</jats:p