17Drought threatens tropical rainforests over seasonal to decadal timescales [1][2][3][4] , but the drivers 18 of tree mortality following drought remain poorly understood 5,6 . It has been suggested that 19 reduced availability of non-structural carbohydrates (NSC) critically increases mortality risk 20 through insufficient carbon supply to metabolism ('carbon starvation') 7,8 . However little is 21 known about how NSC stores are affected by drought, especially over the long term, and 22 whether they are more important than hydraulic processes in determining drought-induced 23 mortality. Using data from the world's longest-running experimental drought study in tropical 24 rainforest (in the Brazilian Amazon), we test whether carbon starvation or deterioration of the 25 water-conducting pathways from soil to leaf trigger tree mortality. Biomass loss from 26 mortality in the experimentally-droughted forest increased substantially after >10 years of 27 reduced soil moisture availability. The mortality signal was dominated by the death of large 28 trees, which were at a much greater risk of hydraulic deterioration than smaller trees. 29However, we find no evidence that the droughted trees suffered carbon starvation, as their 30 NSC concentrations were similar to those of un-droughted trees, and growth rates did not 31 decline in either living or dying individuals. Our results indicate that hydraulics, rather than 32 carbon starvation, triggers tree death from drought in tropical rainforest. 34Drought-response observations from both field-scale experiments and natural droughts have 35 demonstrated increased mortality over the short-term (1-3 years), with notably higher 36 vulnerability for some taxa, and for larger trees 6,9,10 . After several years of drought, 37 recovering growth rates in smaller trees, dbh (diameter at breast height) <40 cm, and reduced 38 mortality have been recorded at different locations 6,11,12 . However, the long-term (>10 yr) 39 sensitivity of tropical forests to predicted prolonged and repeated water deficit [1][2][3] we synthesise these data to test whether long-term soil moisture deficit alters NSC storage 64 and use in tropical rainforest trees, and if this, or hydraulic processes, are most strongly 65 associated with increased mortality rates. 66By 2014, following 13 years of the TFE treatment, cumulative biomass loss through mortality 67 was 41.0±2.7% relative to pre-treatment values (Fig. 1a), and the rate of loss had increased 68 substantially since the previous reported value of 17.2±0.8%, after 7 years of TFE 6 . 69Accelerating biomass loss and failure to recover substantially, or to reach a new 70 equilibrium 13 , has led to a committed flux to the atmosphere from decomposing necromass of 71 101.9±19.1 Mg C ha -1 (Fig. 1a). This biomass loss has been driven by elevated mortality in 72 the largest trees (Fig. 1b), as previously observed over shorter timescales 6 , and has created a 73 canopy that has had a persistently lower average leaf area index during 2010-2014 74 (12.0±1...

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Specific root length (SRL, m g71 ) is probably the most frequently measured morphological parameter of fine roots. It is believed to characterize economic aspects of the root system and to be indicative of environmental changes. The main objectives of this paper were to review and summarize the published SRL data for different tree species throughout Europe and to assess SRL under varying environmental conditions. Meta-analysis was used to summarize the response of SRL to the following manipulated environmental conditions: fertilization, irrigation, elevated temperature, elevated CO 2 , Al-stress, reduced light, heavy metal stress and physical disturbance of soil. SRL was found to be strongly dependent on the fine root classes, i.e. on the ectomycorrhizal short roots (ECM), and on the roots 50.5 mm, 51 mm, 52 mm and 1 -2 mm in diameter SRL was largest for ECM and decreased with increasing diameter. Changes in soil factors influenced most strongly the SRL of ECM and roots 50.5 mm. The variation in the SRL components, root diameter and root tissue density, and their impact on the SRL value were computed. Meta-analyses showed that SRL decreased significantly under fertilization and Al-stress; it responded negatively to reduced light, elevated temperature and CO 2. We suggest that SRL can be used successfully as an indicator of nutrient availability to trees in experimental conditions.

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2. We present a 15-year dataset of litterfall (leaf, flower and fruit, and twigs) from the world's only long-running drought experiment in tropical forest. This dataset comprises one of the longest published litterfall time series in natural forest, which allows the long-term effects of drought on forest reproduction and canopy investment to be explored.3. Over the first 4 years of the experiment, the experimental soil moisture deficit created only a small decline in total litterfall and leaf fall (12% and 13%, respectively), but a very strong initial decline in reproductive litterfall (flowers and fruits) of 54%.This loss of flowering and fruiting was accompanied by a de-coupling of all litterfall patterns from seasonal climate variables. However, following >10 years of the experimental drought, flower and fruiting re-stabilised at levels greater than in the control plot, despite high tree mortality in the drought plot. Litterfall relationships with atmospheric drivers were re-established alongside a strong new apparent trade-off between litterfall and tree growth. 4. Synthesis. We demonstrate that this tropical forest went through an initial shock response during the first 4 years of intense drought, where reproductive effort was arrested and seasonal litterfall patterns were lost. However, following >10 years of experimental drought, this system appears to be re-stabilising at a new functional state where reproduction is substantially elevated on a per tree basis; and there is 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.

Solar irradiance and precipitation are the most likely drivers of the seasonal variation of net primary productivity (NPP) in tropical forests. Since their roles remain poorly understood, we use litter traps, dendrometer bands and census data collected from one hectare permanent plots to quantify the seasonality of above-ground NPP components and weather parameters in 13 sites distributed along a 2800-m altitudinal gradient ranging from lowland Amazonia to the high Andes. We combine canopy leaf area index and litterfall data to describe the seasonality of canopy production. We hypothesize that solar irradiance is the primary driver of canopy phenology in wetter sites, whereas precipitation drives phenology in drier systems. The seasonal rhythm of canopy NPP components is in synchrony with solar irradiance at all altitudes. Leaf litterfall peaks in the late dry season, both in lowland (averaging 0.54 ± 0.08 Mg C ha y −1 , n = 5) and montane forests (averaging 0.29 ± 0.04 Mg C ha y −1 , n = 8). Peaks in above-ground coarse woody NPP appears to be triggered by the onset of rainfall in seasonal lowland rain forests (averaging 0.26 ± 0.04 Mg C ha y −1 , n = 5, in November), but not in montane cloud forests.

Why do some forests produce biomass more efficiently than others? Variations in Carbon Use Efficiency (CUE: total Net Primary Production (NPP)/ Gross Primary Production (GPP)) may be due to changes in wood residence time (Biomass/NPP wood ), temperature, or soil nutrient status. We tested these hypotheses in 14, one ha plots across Amazonian and Andean forests where we measured most key components of net primary production (NPP: wood, fine roots, and leaves) and autotrophic respiration (R a ; wood, rhizosphere, and leaf respiration). We found that lower fertility sites were less efficient at producing biomass and had higher rhizosphere respiration, indicating increased carbon allocation to belowground components. We then compared wood respiration to wood growth and rhizosphere respiration to fine root growth and found that forests with residence times <40 yrs had significantly lower maintenance respiration for both wood and fine roots than forests with residence times >40 yrs. A comparison of rhizosphere respiration to fine root growth showed that rhizosphere growth respiration was significantly greater at low fertility sites. Overall, we found that Amazonian forests produce biomass less efficiently in stands with residence times >40 yrs and in stands with lower fertility, but changes to long-term mean annual temperatures do not impact CUE.

Vegetation processes are fundamentally limited by nutrient and water availability, the uptake of which is mediated by plant roots in terrestrial ecosystems. While tropical forests play a central role in global water, carbon, and nutrient cycling, we know very little about tradeoffs and synergies in root traits that respond to resource scarcity. Tropical trees face a unique set of resource limitations, with rock-derived nutrients and moisture seasonality governing many ecosystem functions, and nutrient versus water availability often separated spatially and temporally. Root traits that characterize biomass, depth distributions, production and phenology, morphology, physiology, chemistry, and symbiotic relationships can be predictive of plants’ capacities to access and acquire nutrients and water, with links to aboveground processes like transpiration, wood productivity, and leaf phenology. In this review, we identify an emerging trend in the literature that tropical fine root biomass and production in surface soils are greatest in infertile or sufficiently moist soils. We also identify interesting paradoxes in tropical forest root responses to changing resources that merit further exploration. For example, specific root length, which typically increases under resource scarcity to expand the volume of soil explored, instead can increase with greater base cation availability, both across natural tropical forest gradients and in fertilization experiments. Also, nutrient additions, rather than reducing mycorrhizal colonization of fine roots as might be expected, increased colonization rates under scenarios of water scarcity in some forests. Efforts to include fine root traits and functions in vegetation models have grown more sophisticated over time, yet there is a disconnect between the emphasis in models characterizing nutrient and water uptake rates and carbon costs versus the emphasis in field experiments on measuring root biomass, production, and morphology in response to changes in resource availability. Closer integration of field and modeling efforts could connect mechanistic investigation of fine-root dynamics to ecosystem-scale understanding of nutrient and water cycling, allowing us to better predict tropical forest-climate feedbacks.