Nutrient stoichiometry and biological dynamics

Banner Image: Common beehive cactus grows in many parts of the American Southwest. Nutrient and water limitation in the desert force many interesting adaptations.All images on this website are by Anshuman Swain, unless stated otherwise. Please seek permission before use.

1. A mathematical model of the Warburg Effect

Project members: Anshuman Swain and William F Fagan

Status: Published; Swain, A. and Fagan, W.F., 2018. A mathematical model of the Warburg Effect: Effects of cell size, shape and substrate availability on growth and metabolism in bacteria. Mathematical biosciences and engineering: MBE, 16(1), pp.168-186.

https://doi.org/10.3934/mbe.2019009

The Warburg effect refers to a curious behavior observed in many organisms and cell types including cancer cells, yeast and bacteria, wherein both the efficient aerobic pathway and the inefficient fermentation pathway are utilized for respiration, despite the presence of ample oxygen. Also termed as overflow metabolism in bacteria, this phenomena has remained an enigmatic and poorly understood phenomenon despite years of experimental work. Here, we focus on bacterial cells and build a model of three trade offs involved in the utilization of aerobic and anaerobic respiration pathways (rate versus yield, surface area versus volume, and fast versus slow biomass production) to explain the observed behavior in cellular systems. The model so constructed also predicts changes in the relative usage of both pathways in terms of size and shape constraints of the cell, and identifies how substrate availability influences growth rate. Additionally, we use the model to explain certain complex phenomena in modern-and paleo-ecosystems, via the concept of overflow metabolism.

A simplified representation of the model. The external glucose is imported into the bacterial call via glucose transporters located on the cell membrane at a rate r0. This glucose is utilized by upper glycolysis pathways to convert it to pyruvate at a rate r1. This converted substrate is then divided into two paths: either to OP (at a rate r2 at the cytochrome oxidase located in the membrane) or to fermentation/aerobic glycolysis (AG) (at a rate r3 at the fermentation enzyme sites located in the cytoplasm). Note that only the substrate used in the latter shunt goes into biomass production whereas substrate used in OP is lost as CO2.

2. Ecological Stoichiometry of the Mountain Cryosphere

Project members: Ze Ren*, Nicolas Martyniuk*, Isabella A. Oleksy*, Anshuman Swain* & Scott Hotaling;

*contributed equally

Status: Published; Ren, Z., Martyniuk, N., Oleksy, I.A., Swain, A. and Hotaling, S., 2019. Ecological stoichiometry of the mountain cryosphere. Frontiers in Ecology and Evolution, 7, p.360.

https://doi.org/10.3389/fevo.2019.00360

Roughly 10% of the Earth's surface is permanently covered by glaciers and ice sheets and in mountain ecosystems, this proportion of ice cover is often even higher. From an ecological perspective, ice-dominated ecosystems place harsh controls on life including cold temperature, limited nutrient availability, and often prolonged darkness due to snow cover for much of the year. Despite these limitations, glaciers, and perennial snowfields support diverse, primarily microbial communities, though macroinvertebrates and vertebrates are also present. The availability and mass balance of key elements [(carbon (C), nitrogen (N), phosphorous (P)] are known to influence the population dynamics of organisms, and ultimately shape the structure and function of ecosystems worldwide. While considerable attention has been devoted to patterns of biodiversity in mountain cryosphere-influenced ecosystems, the ecological stoichiometry of these habitats has received much less attention. Understanding this emerging research arena is particularly pressing in light of the rapid recession of glaciers and perennial snowfields worldwide. In this review, we synthesize existing knowledge of ecological stoichiometry, nutrient availability, and food webs in the mountain cryosphere (specifically glaciers and perennial snowfields). We use this synthesis to develop more general understanding of nutrient origins, distributions, and trophic interactions in these imperiled ecosystems. We focus our efforts on three major habitats: glacier surfaces (supraglacial), the area beneath glaciers (subglacial), and adjacent downstream habitats (i.e., glacier-fed streams and lakes). We compare nutrient availability in these habitats to comparable habitats on continental ice sheets (e.g., Greenland and Antarctica) and show that, in general, nutrient levels are substantially different between the two. We also discuss how ongoing climate warming will alter nutrient and trophic dynamics in mountain glacier-influenced ecosystems. We conclude by highlighting the pressing need for studies to understand spatial and temporal stoichiometric variation in the mountain cryosphere, ideally with direct comparisons to continental ice sheets, before these imperiled habitats vanish completely.

A conceptual diagram of habitats and nutrient dynamics in mountain glacier ecosystems. Major (A–C) and minor (1, 2, 3, 4) components of the mountain cryosphere discussed in this review are shown. Arrows indicate nutrient flow paths, plus signs (+) indicate sources, while red text and minus signs “–” indicate sinks. Yellow boxes with green text represent organisms. Abbreviations include: allo, allochthonous; aut, autochthonous; DOM, dissolved organic matter, Figure is not drawn to scale.

3. Were Neoarchean atmospheric methane hazes and early Paleoproterozoic glaciations driven by the rise of oxygen in surface environments?

Project members: A Swain, A J Kaufman, M Kalinowski and W F Fagan

Status: In Review; preprint link here

In this work, we introduce a microbial-species-interaction based ordinary differential equation model for understanding the conditions under which methane is preferentially released. Methane was the principal greenhouse gas in the Archean and early Proterozoic eons, where its fluctuation in the atmosphere would have played an important role in regulating the climate and the flux of ultraviolet radiation. Exploring certain first-order environmental controls (such as O2 or resource concentration) in the biological methane cycle, and their interactions with various microbial communities, might allow us to deduce and explain the biological, atmospheric and climatic events preserved in the sedimentary rock record between 2.8 and 2.0 billion years ago. Environmental controls on the dynamics of methane cycling may further explain other repetitious events in deep time, as well as the present-day increase in the methane flux to the atmosphere from wetland environments.

We report in this manuscript that the interplay of resource and O2 availability can cause complex cyclic patterns in methane dynamics irrespective of the size any of the (explored) microbial communities and their associated metabolic processes. Although the exact dynamics depend on certain metabolic parameters, but the system behavior in being cyclic or cyclic depend solely on the interplay of resources and oxygen concentration. Based on these model results, we propose that the cyclicity of methane haze events and glacial episodes in the late Archean and early Proterozoic may have been linked to the progressive increase in oceanic and atmospheric O2.

Flow diagram representation of the model system. Here (in blue) A, B, C, F and G are populations of heterotrophic bacteria, methanogens, aerobic methanotrophs, fermenters and anaerobic methanotrophs, respectively. The flux arrows indicate processes that produce (in red) either metabolic reactants or gases cycled among the microbial consortia.