Under the current paradigm, organic matter decomposition and nutrient cycling rates are a function of the imbalance between substrate and microbial biomass stoichiometry. Challenging this view, we demonstrate that in an individual‐based model, microbial community dynamics alter relative C and N limitation during litter decomposition, leading to a system behaviour not predictable from stoichiometric theory alone. Rather, the dynamics of interacting functional groups lead to an adaptation at the community level, which accelerates nitrogen recycling in litter with high initial C : N ratios and thus alleviates microbial N limitation. This mechanism allows microbial decomposers to overcome large imbalances between resource and biomass stoichiometry without the need to decrease carbon use efficiency (CUE), which is in contrast to predictions of traditional stoichiometric mass balance equations. We conclude that identifying and implementing microbial community‐driven mechanisms in biogeochemical models are necessary for accurately predicting terrestrial C fluxes in response to changing environmental conditions.

The supply of nitrogen commonly limits plant production in boreal forests and also affects species composition and ecosystem functions other than plant growth. These interrelations vary across the landscapes, with the highest N availability, plant growth and plant species richness in ground-water discharge areas (GDAs), typically in toe-slope positions, which receive solutes leaching from the much larger groundwater recharge areas (GRAs) uphill. Plant N sources include not only inorganic N, but, as heightened more recently, also organic N species. In general, also the ratio inorganic N over organic N sources increase down hillslopes. Here, we review recent evidence about the nature of the N limitation and its variations in Fennoscandian boreal forests and discuss its implications for forest ecology and management.\ud \ud The rate of litter decomposition has traditionally been seen as the determinant of the rate of N supply. However, while N-rich litter decomposes faster than N-poor litter initially, N-rich litter then decomposes more slowly, which means that the relation between N % of litter and its decomposability is complex. Moreover, in the lower part of the mor-layer, where the most superficial mycorrhizal roots first appear, and N availability matters for plants, the ratio of microbial N over total soil N is remarkably constant over the wide range in litter and soil C/N ratios of between 15 and 40 for N-rich and N-poor sites, respectively. Nitrogen-rich and -poor sites thus differ in the sizes of the total N pool and the microbial N pool, but not in the ratio between them. A more important difference is that the soil microbial N pool turns over faster in N-rich systems because the microbes are more limited by C, while microbes in N-poor systems are a stronger sink for available N.\ud \ud Furthermore, litter decomposition in the most superficial soil horizon (as studied by the so-called litter-bag method) is associated with a dominance of saprotrophic fungi, and absence of mycorrhizal fungi. The focal zone in the context of plant N supply in N-limited forests is further down the soil profile, where ectomycorrhizal (ECM) roots become abundant. Molecular evidence and stable isotope data indicate that in the typical N-poor boreal forests, nitrogen is retained in saprotrophic fungi, likely until they run out of energy (available C-compounds). Then, as heightened by recent research, ECM fungi, which are supplied by photosynthate from the trees, become the superior competitors for N.\ud \ud In N-poor boreal soils strong N retention by microorganisms keeps levels of available N very low. This is exacerbated by an increase in tree C allocation to mycorrhizal fungi (TCAM) relative to net primary production (NPP) with decreasing soil N supply, which causes ECM fungi to retain much of the available soil N for their own growth and transfer little to their tree hosts. The transfer of N through the ECM fungi, and not the rate of litter decomposition, is likely limiting the rate of tree N supply under such conditions. All but a few stress-tolerant less N-demanding plant species, like the ECM trees themselves and ericaceous dwarf shrubs, are excluded.\ud \ud With increasing N supply, a weakening of ECM symbiosis caused by the relative decline in TCAM contributes to shifts in soil microbial community composition from fungal dominance to bacterial dominance. Thus, bacteria, which are less C-demanding, but more likely to release N than fungi, take over. This, and the relatively high pH in GDA, allow autotrophic nitrifying bacteria to compete successfully for the NH4+ released by C-limited organisms and causes the N cycle to open up with leaching of nitrate (NO3−) and gaseous N losses through denitrification. These N-rich conditions allow species-rich communities of N-demanding plant species. Meanwhile, ECM fungi have a smaller biomass, are supplied with N in excess of their demand and will export more N to their host trees. Hence, the gradient from low to high N supply is characterized by profound variations in plant and soil microbial physiologies, especially their relations to the C-to-N supply ratio. We propose how interactions among functional groups can be understood and modelled (the plant-microbe carbon-nitrogen model).\ud \ud With regard to forest management these perspectives explain why the creation of larger tree-free gaps favors the regeneration of tree seedlings under N-limited conditions through reduced belowground competition for N, and why such gaps are less important under high N supply (but when light might be limiting). We also discuss perspectives on the relations between N supply, biodiversity, and eutrophication of boreal forests from N deposition or forest fertilization

We review approaches to predicting carbon and nitrogen allocation in forest models in terms of their underlying assumptions and their resulting strengths and limitations. Empirical and allometric methods are easily developed and computationally efficient, but lack the power of evolution-based approaches to explain and predict multifaceted effects of environmental variability and climate change. In evolution-based methods, allocation is usually determined by maximization of a fitness proxy, either in a fixed environment, which we call optimal response (OR) models, or including the feedback of an individual's strategy on its environment (game-theoretical optimization, GTO). Optimal response models can predict allocation in single trees and stands when there is significant competition only for one resource. Game-theoretical optimization can be used to account for additional dimensions of competition, e.g., when strong root competition boosts root allocation at the expense of wood production. However, we demonstrate that an OR model predicts similar allocation to a GTO model under the root-competitive conditions reported in free-air carbon dioxide enrichment (FACE) experiments. The most evolutionarily realistic approach is adaptive dynamics (AD) where the allocation strategy arises from eco-evolutionary dynamics of populations instead of a fitness proxy. We also discuss emerging entropy-based approaches that offer an alternative thermodynamic perspective on allocation, in which fitness proxies are replaced by entropy or entropy production. To help develop allocation models further, the value of wide-ranging datasets, such as FLUXNET, could be greatly enhanced by ancillary measurements of driving variables, such as water and soil nitrogen availability.

Summary: Symbioses between plant roots and mycorrhizal fungi are thought to enhance plant uptake of nutrients through a favourable exchange for photosynthates. Ectomycorrhizal fungi are considered to play this vital role for trees in nitrogen (N)‐limited boreal forests. We followed symbiotic carbon (C)–N exchange in a large‐scale boreal pine forest experiment by tracing ¹³CO₂ absorbed through tree photosynthesis and ¹⁵N injected into a soil layer in which ectomycorrhizal fungi dominate the microbial community. We detected little ¹⁵N in tree canopies, but high levels in soil microbes and in mycorrhizal root tips, illustrating effective soil N immobilization, especially in late summer, when tree belowground C allocation was high. Additions of N fertilizer to the soil before labelling shifted the incorporation of ¹⁵N from soil microbes and root tips to tree foliage. These results were tested in a model for C–N exchange between trees and mycorrhizal fungi, suggesting that ectomycorrhizal fungi transfer small fractions of absorbed N to trees under N‐limited conditions, but larger fractions if more N is available. We suggest that greater allocation of C from trees to ectomycorrhizal fungi increases N retention in soil mycelium, driving boreal forests towards more severe N limitation at low N supply.

Plants respond to nitrogen availability by changing their root : shoot ratios. One hypothesis used to explain this allocation is that plants optimize their behaviour by maximizing their relative growth rate. The consequences of this hypothesis were investigated by formulating two models for root : shoot allocation, with and without explicit inclusion of maintenance respiration. The models also took into account that relative growth rate is a linear function of plant nitrogen concentration. The model without respiration gave qualitatively reasonable results when predictions were compared with observed results from growth experiments with birch and tomato. The explicit inclusion of maintenance respiration improved considerably the agreement between prediction and observation, and for birch was within the experimental accuracy. Further improvements will require additional details in the description of respiratory processes and the nitrogen uptake function. Plants growing under extreme nutrient stress may also optimize their behaviour with respect to other variables in addition to relative growth rate.