ALBERT

Description: Autotrophic organisms reveal an astounding flexibility in their elemental stoichiometry, with potentially major implications on biogeochemical cycles and ecological functioning. Notwithstanding, stoichiometric regulation, and co-limitation by multiple resources in autotrophs were in the past often described by heuristic formulations. In this study, we present a mechanistic model of autotroph growth, which features two major improvements over the existing schemes. First, we introduce the concept of metabolic network independence that defines the degree of phase-locking between accessory machines. Network independence is in particular suggested to be proportional to protein synthesis capability as quantified by variable intracellular N:C. Consequently, the degree of co-limitation becomes variable, contrasting with the dichotomous debate on the use of Liebig's law or the product rule, standing for constantly low and high co-limitation, respectively. Second, we resolve dynamic protein partitioning to light harvesting, carboxylation processes, and to an arbitrary number of nutrient acquisition machineries, as well as instantaneous activity regulation of nutrient uptake. For all regulatory processes we assume growth rate optimality, here extended by an explicit consideration of indirect feed-back effects. The combination of network independence and optimal regulation displays unprecedented skill in reproducing rich stoichiometric patterns collected from a large number of published chemostat experiments. This high skill indicates (1) that the current paradigm of fixed co-limitation is a critical short-coming of conventional models, and (2) that stoichiometric flexibility in autotrophs possibly reflects an optimality strategy. Numerical experiments furthermore show that regulatory mechanisms homogenize the effect of multiple stressors. Extended optimality alleviates the effect of the most limiting resource(s) while down-regulating machineries for the less limiting ones, which induces an ubiquitous response surface of growth rate over ambient resource levels. Our approach constitutes a basis for improved mechanistic understanding and modeling of acclimative processes in autotrophic organisms. It hence may serve future experimental and theoretical investigations on the role of those processes in aquatic and terrestrial ecosystems.

Description: Field data collected for the North Sea indicate a prominent seasonal variation in the vertical distribution of total organic carbon (TOC) and macrobenthic biomass in sediments. The vertical TOC profiles classify into three modes, with maximum at surface, middle and deep part of sediments, respectively. We here present a mechanistic model to quantify, for the first time, the dynamic interaction between sedimentary TOC and benthic fauna. The major model principles include that (i) the vertical distribution of macrobenthic biomass is a trade-off between nutritional benefit (quantity and quality of TOC) and the costs of burial (respiration) and mortality, and (ii) the vertical transport of TOC is in turn modulated by macrobenthos through bioturbation. A novelty of our model is that bioturbation is resolved dynamically depending on variation of local food resources and macrobenthic biomass. This allows capturing of the benthic response to both depositional and erosional conditions and improving estimates of the material exchange flux at the sediment-water interface. The coupling of the TOC-benthos model with 3D hydrodynamic-ecological simulations reveals that the three profile modes of sedimentary TOC (in both quantify and quality) can be explained as a combined response to pelagic conditions (shear stress and primary production) and the synergy between bioturbation, vertical redistribution of higher quality TOC and vertical positioning of benthic organisms. A model reconstruction of the benthic status in the North Sea from 1950s to 2010s indicates that despite a relatively stable pattern at decadal and regional scales, significant variations exist at smaller scales characterized by seasons and local areas. In addition, inter-annual and multi-year cycle-like variations are also prominent especially in coastal areas.