Exploration capacity versus specific enzymatic activity of ectomycorrhizas in response to primary productivity and soil phosphorus availability in Bornean tropical rainforests.

Ectomycorrhizal (ECM) fungi are functionally important in biogeochemical cycles in tropical ecosystems. Extracellular enzymatic activity of ECM on a ground-area basis is the product of two attributes; exploration capacity (ECM surface-area) and specific enzymatic activity. Here, we elucidated which attribute better explained the ECM enzymatic activity in response to different levels of soil phosphorus (P) and Nitrogen (N) availability in five Bornean tropical rainforests. We determined the surface area of ECM root tips as well as the enzymatic activities per ECM surface area for carbon (C), N and P degrading enzymes in each site. We evaluated the relationship of ECM enzyme activities with the resource availabilities of C (Above-ground net primary production; ANPP), N, and P of ECM by a generalized linear mixed model. The ECM enzymatic activities on a ground-area basis were more significantly determined by specific enzymatic activity than by the exploration capacity. Specific enzymatic activities were generally negatively affected by C (ANPP) and soil P availability. ECM fungi enhance the specific enzyme activity rather than the exploration capacity to maintain the capacity of nutrient acquisition. The less dependence of ECM fungi on the exploration capacity in these forests may be related to the limitation of C supply from host trees. We highlighted the adaptive mechanisms of ECM fungi on nutrient acquisition in tropical ecosystems through the response of enzymatic activity to nutrient availability across the elements.

Roles of lower-side leaf trichomes in diffusion resistance and gas-exchange characteristics across environmental gradients in Metrosideros polymorpha.

Leaf trichomes on the lower leaf surface are common in many plant species, especially those grown under dry and/or low-temperature conditions; however, their adaptive significance remains unclear. Lower-side leaf trichomes can directly decrease gas fluxes through increased gas-diffusion resistance but can indirectly increase gas fluxes through increased leaf temperature owing to increased heat-diffusion resistance. We examined whether the combined direct and indirect effects of trichome resistance increase photosynthetic rates and water-use efficiency (WUE) using Metrosideros polymorpha Gaud., which varies widely in the masses of lower-side non-glandular leaf trichomes across various environments on the Hawaiian Islands. We employed both field surveys, including ecophysiological measurements at five elevation sites, and simulation analyses to predict the gas-exchange rates of leaves with various trichome-layer thicknesses across a wide range of environmental conditions. Field surveys showed that the trichome-layer thickness was the largest at the coldest and driest site and the thinnest at the wettest site. Field surveys, experimental manipulations and simulation analyses demonstrated that leaf trichomes significantly increased leaf temperature owing to the increased heat resistance. Simulation analyses showed that the effect of leaf trichomes on heat resistance was much larger than that on gas-flux resistance. Leaf trichomes can increase daily photosynthesis only in cold dry areas by increasing the leaf temperature. However, the increased leaf temperature with leaf trichomes resulted in a consistent decrease in the daily WUE at all elevation sites. The magnitudes of trichome effects on gas-exchange rates were associated with the temperature difference across the elevational gradient, the strong light intensity in Hawaii, the leaf-size variation and the conservative stomatal behavior of M. polymorpha as well as the trichome-layer thickness. In summary, the lower-side leaf trichomes in M. polymorpha can be beneficial for carbon assimilation in low-temperature environments but not for water conservation in most environments in terms of diffusion resistance.

Interspecific differences in the responses of root phosphatase activities and morphology to nitrogen and phosphorus fertilization in Bornean tropical rain forests.

Soil organic phosphorus (P) compounds can be the main P source for plants in P-limited tropical rainforests. Phosphorus occurs in diverse chemical forms, including monoester P, diester P, and phytate, which require enzymatic hydrolysis by phosphatase into inorganic P before assimilation by plants. The interactions between plant interspecific differences in organic P acquisition strategies via phosphatase activities with root morphological traits would lead to P resource partitioning, but they have not been rigorously evaluated. We measured the activities of three classes of phosphatases (phosphomonoesterase, PME; phosphodiesterase, PDE; and phytase, PhT), specific root length (SRL), root diameter, and root tissue density in mature tree species with different mycorrhizal associations (ectomycorrhizal [ECM] or arbuscular mycorrhizal [AM]) and different successional status (climax or pioneer species) in Sabah, Malaysia. We studied nitrogen (N)- and P-fertilized plots to evaluate the acquisition strategies for organic P under P-limited conditions 7 years after fertilization was initiated. P fertilization reduced the PME activity in all studied species and reduced PhT and PDE activities more in climax species than in the two pioneer species, irrespective of the mycorrhizal type. PDE activity increased in some climax species after N fertilization, suggesting that these species allocate excess N to the synthesis of PDE. Moreover, PME and PhT activities, but not PDE activity, correlated positively with SRL. We suggest that climax species tend to be more strongly dependent on recalcitrant organic P (i.e., phytate and/or diester P) than pioneer species, regardless of the mycorrhizal type. We also suggest that trees in which root PME or PhT activity is enhanced can increase their SRL to acquire P efficiently. Resource partitioning of soil organic P would occur among species through differences in their phosphatase activities, which plays potentially ecologically important role in reducing the competition among coexisting tree species in lowland tropical rainforests.

A cost-benefit analysis of leaf carbon economy with consideration of seasonal changes in leaf traits for sympatric deciduous and evergreen congeners: implications for their coexistence.

Deciduous and evergreen species, which have evolved repeatedly across different clades, can coexist in a given environment despite substantial differences in their leaf traits. It remains unclear how these two groups differ in the development of leaf traits over their lifespans or how their carbon economy - the balance between lifetime carbon gain and leaf construction cost - is determined. We determined the photosynthetic rate (Aarea ), leaf mass per area (LMA), leaf mechanical strength and leaf water potentials and estimated the lifetime carbon gain and leaf construction cost of five closely related pairs of evergreen and deciduous species co-occurring in a temperate forest. Aarea of evergreen species was lower during their first spring, similar in summer and higher than the autumn until the following spring than their deciduous counterparts. Leaf mechanical strength, osmotic pressures and LMA increased continuously towards winter in evergreen species while remaining largely constant in deciduous species. The ratio of lifetime carbon gain to leaf construction cost was similar between the two groups. The additional cost associated with enduring winter is paid back by a longer revenue of photosynthesis in evergreen species, allowing evergreen and deciduous leaf habits to coexist in the seasonal environment.

Experimental Amelioration of Harsh Weather Speeds Growth and Development in a Tropical Montane Songbird.

AbstractOrganisms living at high elevations generally grow and develop more slowly than those at lower elevations. Slow montane ontogeny is thought to be an evolved adaptation to harsh environments that improves juvenile quality via physiological trade-offs. However, slower montane ontogeny may also reflect proximate influences of harsh weather on parental care and offspring development. We experimentally heated and protected nests from rain to ameliorate harsh montane weather conditions for mountain blackeyes (Chlorocharis emiliae), a montane songbird living at approximately 3,200 m asl in Malaysian Borneo. This experiment was designed to test whether cold and wet montane conditions contribute to parental care and postnatal growth and development rates at high elevations. We found that parents increased provisioning and reduced time spent warming offspring, which grew faster and departed the nest earlier compared with offspring from unmanipulated nests. Earlier departure reduces time-dependent predation risk, benefitting parents and offspring. These plastic responses highlight the importance of proximate weather contributions to broad patterns of montane ontogeny and parental care.

Light and nutrient limitations for tree growth on young versus old soils in a Bornean tropical montane forest.

We examined forest and tree responses to decreasing nutrient availability with soil aging in a species-rich tropical montane rain forest on Mount Kinabalu, Borneo. Community composition and structure and tree growth rates were compared between two 1 ha plots on nutrient-rich young soil versus nutrient-deficient old soil. Myrtaceae and Fagaceae dominated both plots. With soil aging, the dominance of Lauraceae, stem density, basal area and aboveground biomass decreased, and the forest understory became brighter. Some dominant taxa on the old soil (Podocarpaceae and the genus Tristaniopsis in Myrtaceae) were virtually absent on the young soil; this was attributed to light limitation in the understory. Growth rates of understory trees were lower on the young soil, whereas those of canopy trees were lower on the old soil. This suggested that the growth of understory trees was limited by light on the young soil, whereas that of canopy trees was limited by nutrients on the old soil. Of the eight species that were abundant in both plots, the dominance of five species was considerably lower on the old soil, four of which also exhibited decreased maximum sizes and lower growth rates. The remaining three species showed similar dominance across plots without a decline in growth rates, although they exhibited decreased maximum sizes on the old soil. These analyses demonstrated divergent responses of species to the soil-age gradient. We suggest that the differential responses of species to decreasing nutrient availability with a concomitant increase in understory light levels explain floristic turnover with soil aging.

Leaf trichomes in Metrosideros polymorpha can contribute to avoiding extra water stress by impeding gall formation.

BACKGROUND AND AIMS: Plants inhabiting arid environments tend to have leaf trichomes, but their adaptive significance remains unclear. Leaf trichomes are known to play a role in plant defence against herbivores, including gall makers. Because gall formation can increase water loss partly through increased surface area, we tested the novel hypothesis that leaf trichomes could contribute to avoiding extra water stress by impeding gall formation, which would have adaptive advantages in arid environments. METHODS: We focused on Metrosideros polymorpha, an endemic tree species in the Hawaiian Islands, whose leaves often suffer from galls formed by specialist insects, Hawaiian psyllids (Pariaconus spp.). There is large variation in the amount of leaf trichomes (0-40 % of leaf mass) in M. polymorpha. Three gall types are found on the island of Hawaii: the largest is the 'cone' type, followed by 'flat' and 'pit' types. We conducted laboratory experiments to quantify the extent to which gall formation is associated with leaf water relations. We also conducted a field census of 1779 individuals from 48 populations across the entire range of habitats of M. polymorpha on the island of Hawaii to evaluate associations between gall formation (presence and abundance) and the amount of leaf trichomes. KEY RESULTS: Our laboratory experiment showed that leaf minimum conductance was significantly higher in leaves with a greater number of cone- or flat-type galls but not pit-type galls. Our field census suggested that the amount of trichomes was negatively associated with probabilities of the presence of cone- or flat-type galls but not pit-type galls, irrespective of environmental factors. CONCLUSION: Our results suggest that leaf trichomes in M. polymorpha can contribute to the avoidance of extra water stress through interactions with some gall-making species, and potentially increase the fitness of plants under arid conditions.

Meta-analysis of management effects on biodiversity in plantation and secondary forests of Japan.

Conservation of temperate forest biodiversity has historically focused on natural old-growth. Less than 3% of the world's temperate forests remain unmodified by humans, however, and much of temperate-forest biodiversity is held in the predominating planted and secondary forests. Japan provides a widely applicable model for examining how to maximize biodiversity in managed temperate forests, because of its richness of forestry research generated from its vast forest area, albeit largely in Japanese, and the wide practice of its dominant management interventions across the northern temperate zone. Management for plantations includes thinning, extended rotation cycles and clear-cutting. For secondary forests regenerating from past clearance, traditional management varies in its intensities, from clear-cutting as coppices to small-scale understory clearance. Here we provide a first synthesis of published research on biodiversity in planted and secondary forests of Japan, relevant to management of these types of forest in northern temperate regions. Systematic review and meta-analyses of papers published in English and Japanese quantified management impacts on species richness and abundance of several taxa, in relation to moderator variables including stand age and management intensity. Plantation thinning substantially increases the richness and abundance of several taxa. Effect sizes decline with time since thinning for the abundance of regenerating saplings and seedlings, necessitating repeated thinning treatments every 6 years to sustain this positive effect. Taxonomic groups exhibit variable relationships with stand age in both planted and secondary forests, indicating a need to include both young and old forest stands in managed forest mosaics. We find an insufficient evidence base is available to allow for a meaningful synthesis of low-intensity management effects in historically managed secondary forests, with studies varying widely in scale and reported outcomes. We outline an agenda for the research community to achieve a systematic evaluation of scale-dependent effects of traditional forest management on biodiversity.

Author Correction: Long-term carbon sink in Borneo's forests halted by drought and vulnerable to edges.

The original version of this Article contained an error in the third sentence of the abstract and incorrectly read "Here, using long-term plot monitoring records of up to half a century, we find that intact forests in Borneo gained 0.43 Mg C ha-1 year-1 (95% CI 0.14-0.72, mean period 1988-2010) above-ground live biomass", rather than the correct "Here, using long-term plot monitoring records of up to half a century, we find that intact forests in Borneo gained 0.43 Mg C ha-1 year-1 (95% CI 0.14-0.72, mean period 1988-2010) in above-ground live biomass carbon". This has now been corrected in both the PDF and HTML versions of the Article.

Long-term carbon sink in Borneo's forests halted by drought and vulnerable to edge effects.

Less than half of anthropogenic carbon dioxide emissions remain in the atmosphere. While carbon balance models imply large carbon uptake in tropical forests, direct on-the-ground observations are still lacking in Southeast Asia. Here, using long-term plot monitoring records of up to half a century, we find that intact forests in Borneo gained 0.43 Mg C ha-1 per year (95% CI 0.14-0.72, mean period 1988-2010) above-ground live biomass. These results closely match those from African and Amazonian plot networks, suggesting that the world's remaining intact tropical forests are now en masse out-of-equilibrium. Although both pan-tropical and long-term, the sink in remaining intact forests appears vulnerable to climate and land use changes. Across Borneo the 1997-1998 El Niño drought temporarily halted the carbon sink by increasing tree mortality, while fragmentation persistently offset the sink and turned many edge-affected forests into a carbon source to the atmosphere.

Phosphorus and nitrogen resorption from different chemical fractions in senescing leaves of tropical tree species on Mount Kinabalu, Borneo.

Nutrient resorption, a process by which plants degrade organic compounds and resorb their nutrients from senescing tissues, is a crucial plant function to increase growth and fitness in nutrient-poor environments. Tropical trees on phosphorus (P)-poor soils are particularly known to have high P-resorption efficiency (PRE, the percentage of P resorbed from senescing leaves before abscission per total P in green leaves). However, the biochemical mechanisms underlying this greater PRE remain unclear. In this study, we determined the P concentration in easily soluble, nucleic acid, lipid and residual fractions for green and senescent leaves of 22 tree species from three sites, which differed in P availability, on the lower flanks of Mt. Kinabalu, Borneo. PRE varied from 24 to 93% and was higher in species from the P-poor site. P-resorption rate was greatest from the lipid fraction, the nucleic acid fraction, and lowest in the easily soluble fraction and the residual fraction when all the species were pooled. For species with higher PRE, P-resorption rate of the residual fraction was relatively high and was comparable in magnitude to that of the other labile fractions. This suggests that tree species inhabiting P-poor environments increased PRE by improving the degradation of recalcitrant compounds. This study suggests that plants selectively degrade organic compounds depending on environmental conditions, which is a key mechanism underlying the variation of PRE.

The population genomic signature of environmental association and gene flow in an ecologically divergent tree species Metrosideros polymorpha (Myrtaceae).

Genomewide markers enable us to study genetic differentiation within a species and the factors underlying it at a much higher resolution than before, which advances our understanding of adaptation in organisms. We investigated genomic divergence in Metrosideros polymorpha, a woody species that occupies a wide range of ecological habitats across the Hawaiian Islands and shows remarkable phenotypic variation. Using 1659 single nucleotide polymorphism (SNP) markers annotated with the genome assembly, we examined the population genetic structure and demographic history of nine populations across five elevations and two ages of substrates on Mauna Loa, the island of Hawaii. The nine populations were differentiated into two genetic clusters distributed on the lower and higher elevations and were largely admixed on the middle elevation. Demographic modelling revealed that the two genetic clusters have been maintained in the face of gene flow, and the effective population size of the high-altitude cluster was much smaller. A FST -based outlier search among the 1659 SNPs revealed that 34 SNPs (2.05%) were likely to be under divergent selection and the allele frequencies of 21 of them were associated with environmental changes along elevations, such as temperature and precipitation. This study shows a genomic mosaic of M. polymorpha, in which contrasting divergence patterns were found. While most genomic polymorphisms were shared among populations, a small fraction of the genome was significantly differentiated between populations in diverse environments and could be responsible for the dramatic adaptation to a wide range of environments.

Contrasting effects of exogenous phosphorus application on N2O emissions from two tropical forest soils with contrasting phosphorus availability.

An incubation study was conducted to test the effects of phosphorus (P) addition on nitrous oxide (N2O) emissions from the soils taken from two tropical rain forests established on different parent materials [meta-sedimentary (MS) and ultrabasic (UB) rock] on Mt. Kinabalu, Borneo. Earlier studies suggest that the forest on UB soils is more strongly limited by P than that on MS soils is. In MS soils, P addition significantly reduced N2O emissions. Since neither ammonium (NH4 (+)) nor nitrate (NO3 (-)) contents were reduced by P addition, we assumed that the decrease in N2O emissions were not due to the previously-reported mechanism: P addition stimulated microbial nitrogen (N) immobilization and collateral inorganic N consumption, reducing resources for producing N2O. Since P addition enhanced the ratios of microbial biomass to CO2 and N2O emissions (indicators of nitrifying and/or denitrifying respiratory efficiency), it was suggested that the N required for the respiration of nitrifying and/or denitrifying bacteria was reduced, leading to reduced N2O emissions. On the other hand, P addition had no effects on N2O emissions in UB soils. The respiratory efficiency did not change significantly by P addition, possibly because the microbial community in the highly-P-depleted UB soils shifted by P addition, with which the enhancement of respiration efficiency did not co-vary. We concluded that (1) P addition may control N2O emissions through increasing respiratory efficiency, and (2) the effects may be different depending on the differences in P availability.

Nutrient allocation among plant organs across 13 tree species in three Bornean rain forests with contrasting nutrient availabilities.

Allocation of nitrogen (N) and phosphorus (P) among plant organs is an important factor regulating growth rate, which is a key ecological process associated with plant life-history strategies. However, few studies have explored how N and P investment in photosynthetic (leaves) and non-photosynthetic (stems and roots) organs changes in relation to depletion of each element. We investigated nutrient concentrations of plant organs in relation to whole-plant nutrient concentration (total nutrient weight per total biomass) as an index of nutrient status of each individual using the saplings of the 13 species in three tropical rain forests with contrasting N and P availabilities (tropical evergreen forests and tropical heath forests). We found a steeper decrease in foliar N concentration than foliar P concentration with decreasing whole-plant nutrient concentration. Moreover, the steeper decrease in foliar N concentration was associated with relatively stable N concentration in stems, and vice versa for P. We suggest that the depletion of N is associated with a rapid dilution of foliar N because the cell walls in non-photosynthetic organs function as an N sink. On the other hand, these species can maintain foliar P concentration by decreasing stem P concentrations despites the depletion of P. Our results emphasize the significance of non-photosynthetic organs as an N sink for understanding the variation of foliar nutrient concentrations for the tree species in the three Bornean rain forests with different N and P availabilities.

A quantitative analysis of phenotypic variations of Metrosideros polymorpha within and across populations along environmental gradients on Mauna Loa, Hawaii.

Metrosideros polymorpha, a dominant tree species in the Hawaiian Islands, shows an extreme phenotypic polymorphism both across gradients of climatic/edaphic conditions and within populations, making it a potentially useful model species for evolutionary study. In order to understand how the phenotypic diversity is maintained within populations as well as across populations, we examined the diversities of several leaf and stem functional traits across five elevations and two soil substrates on the volcanic mountain of Mauna Loa, on the island of Hawaii. Leaf dry mass per area (LMA), a key leaf functional trait, was particularly focused on and analyzed in relation to its underlying components-namely, tissue LMA and trichome LMA (LMA = tissue LMA + trichome LMA). Across populations, tissue LMA increased linearly with elevation while trichome LMA showed unimodal patterns with elevation, which were better correlated with temperature and rainfall, respectively. Substantial phenotypic variations were also found within populations. Interestingly, the variations of tissue LMA were often negatively correlated to trichome LMA within populations, which contrasts with the cross-populations pattern, where a strong positive correlation between tissue LMA and trichome LMA was found. This suggests that phenotypic variations within populations were substantially influenced by local ecological processes. Soil depth (an indicator of local water availability) and tree size (an indicator of colonized timing) modestly explained the within-population variations, implying other local environmental factors and/or random processes are also important in local phenotypic diversity. This study provides an insight about how phenotypic diversity of plant species is maintained from local to landscape levels.

Relationship between photosynthetic phosphorus-use efficiency and foliar phosphorus fractions in tropical tree species.

How plants develop adaptive strategies to efficiently use nutrients on infertile soils is an important topic in plant ecology. It has been suggested that, with decreasing phosphorus (P) availability, plants increase photosynthetic P-use efficiency (PPUE) (i.e., the ratio of instantaneous photosynthetic carbon assimilation rate per unit foliar P). However, the mechanism to increase PPUE remains unclear. In this study, we tested whether high PPUE is explained by an optimized allocation of P in cells among P-containing biochemical compounds (i.e., foliar P fractions). We investigated the relationships among mass-based photosynthetic carbon assimilation rate (A mass), PPUE, total foliar P concentration, and foliar P fractions in 10 tree species in two tropical montane rain forests with differing soil P availability (five species on sedimentary soils and five species on P-poorer ultrabasic serpentine soils) on Mount Kinabalu, Borneo. We chemically fractionated foliar P into the following four fractions: metabolic P, lipid P, nucleic acid P, and residual P. A mass was positively correlated with the concentrations of total foliar P and of metabolic P across 10 tree species. Mean A mass and mean concentrations of total foliar P and of each foliar P fraction were lower on the P-poorer ultrabasic serpentine soils than on the sedimentary soils. There was a negative relationship between the proportion of metabolic P per total P and the proportion of lipid P per total P. PPUE was positively correlated with the ratio of metabolic P to lipid P. High PPUE is explained by the net effect of a relatively greater investment of P into P-containing metabolites and a relatively lesser investment into phospholipids in addition to generally reduced concentrations of all P fractions. We conclude that plants optimize the allocation of P among foliar P fractions for maintaining their productivity and growth and for reducing demand for P as their adaptation to P-poor soils.

Development of SSR markers for the tropical alpine tree species Leptospermum recurvum (Myrtaceae) on Mount Kinabalu in Borneo.

PREMISE OF THE STUDY: Nuclear microsatellite (simple sequence repeat [SSR]) markers were developed for the woody species Leptospermum recurvum found on Mount Kinabalu, Borneo, to facilitate investigation of the genetic structure and patterns of gene flow in relation to leaf phenotypic polymorphisms. • METHODS AND RESULTS: Eleven primer pairs were developed using the compound SSR marker technique. Ten of the 11 loci were polymorphic and their expected heterozygosity ranged from 0.04 to 0.83. Neither linkage disequilibrium nor departure from Hardy-Weinberg equilibrium were detected. All primer pairs also amplified the SSR loci of L. polygalifolium. • CONCLUSIONS: These findings suggest the utility of these primers for investigating genetic structure and gene flow in L. recurvum and indicate their applicability to another species of Leptospermum.

Co-benefits of sustainable forest management in biodiversity conservation and carbon sequestration.

BACKGROUND: Sustainable forest management (SFM), which has been recently introduced to tropical natural production forests, is beneficial in maintaining timber resources, but information about the co-benefits for biodiversity conservation and carbon sequestration is currently lacking. METHODOLOGY/PRINCIPAL FINDINGS: We estimated the diversity of medium to large-bodied forest-dwelling vertebrates using a heat-sensor camera trapping system and the amount of above-ground, fine-roots, and soil organic carbon by a combination of ground surveys and aerial-imagery interpretations. This research was undertaken both in SFM applied as well as conventionally logged production forests in Sabah, Malaysian Borneo. Our carbon estimation revealed that the application of SFM resulted in a net gain of 54 Mg C ha(-1) on a landscape scale. Overall vertebrate diversity was greater in the SFM applied forest than in the conventionally logged forest. Specifically, several vertebrate species (6 out of recorded 36 species) showed higher frequency in the SFM applied forest than in the conventionally logged forest. CONCLUSIONS/SIGNIFICANCE: The application of SFM to degraded natural production forests could result in greater diversity and abundance of vertebrate species as well as increasing carbon storage in the tropical rain forest ecosystems.

Phenolic control of plant nitrogen acquisition through the inhibition of soil microbial decomposition processes: a plant-microbe competition model.

Phenolics are a dominant class of plant secondary metabolites that have strong effects on various ecosystem processes. The ecological significance of these compounds, however, is still poorly understood. We hypothesized that the inhibitory effects of phenolics on microbial activity could enhance plant nitrogen acquisition by relaxing competition between plants and microbes. To test this hypothesis theoretically, we constructed a novel and simple mechanistic model by unifying two concepts: one is a new paradigm of nitrogen cycling which considers the uptake of organic nitrogen by plants, and the other is that phenolics can regulate nitrogen cycling by inhibiting microbial decomposition processes. Our plant-microbe competition model consists of five compartments (plants, soil microbes, debris, organic nitrogen and inorganic nitrogen) and incorporates the essential processes of nitrogen cycling: plant uptake of monomers, competition between plants and microbes, and the depolymerization process. Our analysis showed that plant nitrogen acquisition was maximized at intermediate levels of phenolics, but only when plants could utilize organic nitrogen. Furthermore, this pattern occurred over a broad range of parameter conditions. Our study successfully demonstrated the potential role of phenolics in plant nitrogen acquisition throughout natural environments.