Environment-dependent metabolic investments in the mixotrophic chrysophyte Ochromonas.

Mixotrophic protists combine photosynthesis and phagotrophy to obtain energy and nutrients. Because mixotrophs can act as either primary producers or consumers, they have a complex role in marine food webs and biogeochemical cycles. Many mixotrophs are also phenotypically plastic and can adjust their metabolic investments in response to resource availability. Thus, a single species's ecological role may vary with environmental conditions. Here, we quantified how light and food availability impacted the growth rates, energy acquisition rates, and metabolic investment strategies of eight strains of the mixotrophic chrysophyte, Ochromonas. All eight Ochromonas strains photoacclimated by decreasing chlorophyll content as light intensity increased. Some strains were obligate phototrophs that required light for growth, while other strains showed stronger metabolic responses to prey availability. When prey availability was high, all eight strains exhibited accelerated growth rates and decreased their investments in both photosynthesis and phagotrophy. Photosynthesis and phagotrophy generally produced additive benefits: In low-prey environments, Ochromonas growth rates increased to maximum, light-saturated rates with increasing light but increased further with the addition of abundant bacterial prey. The additive benefits observed between photosynthesis and phagotrophy in Ochromonas suggest that the two metabolic modes provide nonsubstitutable resources, which may explain why a tradeoff between phagotrophic and phototrophic investments emerged in some but not all strains.

Evidence for evolutionary adaptation of mixotrophic nanoflagellates to warmer temperatures.

Mixotrophs, organisms that combine photosynthesis and heterotrophy to gain energy, play an important role in global biogeochemical cycles. Metabolic theory predicts that mixotrophs will become more heterotrophic with rising temperatures, potentially creating a positive feedback loop that accelerates carbon dioxide accumulation in the atmosphere. Studies testing this theory have focused on phenotypically plastic (short-term, non-evolutionary) thermal responses of mixotrophs. However, as small organisms with short generation times and large population sizes, mixotrophs may rapidly evolve in response to climate change. Here, we present data from a 3-year experiment quantifying the evolutionary response of two mixotrophic nanoflagellates to temperature. We found evidence for adaptive evolution (increased growth rates in evolved relative to acclimated lineages) in the obligately phototrophic strain, but not in the facultative phototroph. All lineages showed trends of increased carbon use efficiency, flattening of thermal reaction norms, and a return to homeostatic gene expression. Generally, mixotrophs evolved reduced photosynthesis and higher grazing with increased temperatures, suggesting that evolution may act to exacerbate mixotrophs' effects on global carbon cycling.

Prey type constrains growth and photosynthetic capacity of the kleptoplastidic ciliate Mesodinium chamaeleon (Ciliophora).

Kleptoplastidic, or chloroplast-stealing, lineages offer insight into the process of acquiring photosynthesis. By quantifying the ability of these organisms to retain and use photosynthetic machinery from their prey, we can understand how intermediaries on the endosymbiosis pathway might have evolved regulatory and maintenance mechanisms. Here, we focus on a mixotrophic kleptoplastidic ciliate, Mesodinium chamaeleon, noteworthy for its ability to retain functional chloroplasts from at least half a dozen cryptophyte algal genera. We contrasted the performance of kleptoplastids from blue-green and red cryptophyte prey as a function of light level and feeding history. Our experiments showed that starved M. chamaeleon cells are able to maintain photosynthetic function for at least 2 weeks and that M. chamaeleon containing red plastids lost chlorophyll and electron transport capacity faster than those containing blue-green plastids. However, likely due to increased pigment content and photosynthetic rates in red plastids, M. chamaeleon had higher growth rates and more prolonged growth when feeding on red cryptophytes. For example, M. chamaeleon grew rapidly and extensively when fed the blue-green cryptophyte Chroomonas mesostigmatica, but this growth appeared to hinge on high levels of feeding supporting photosynthetic activity. In contrast, even starved M. chamaeleon containing red plastids from Rhodomonas salina could achieve high photosynthetic rates and extensive growth. Our findings show that plastid origin impacts the maintenance and magnitude of photosynthetic activity, though whether this is due to variation in ciliate control or gradual loss of plastid function in ingested prey cells remains unknown.

Soil fungal community composition and functional similarity shift across distinct climatic conditions.

A large part of ecosystem function in woodland systems depends on soil fungal communities. However, global climate change has the potential to fundamentally alter these communities as fungal species are filtered with changing environmental conditions. In this study, we examined the potential effects of climate on host-associated (i.e. tree-associated) soil fungal communities at climatically distinct sites in the Tehachapi Mountains in California, where more arid conditions represent likely regional climate futures. We found that soil fungal community composition changes strongly across sites, with species richness and diversity being highest at the most arid site. However, host association may buffer the effects of climate on community composition, as host-associated fungal communities are more similar to each other across climatically distinct sites than the whole fungal community. Lastly, an examination of functional traits for ectomycorrhizal fungi, a well-studied guild of fungal mutualist species, showed that stress-tolerant traits were more abundant at arid sites than mesic sites, providing a mechanistic understanding of these community patterns. Taken together, our results indicate that fungal community composition will likely shift with future climate change but that host association may buffer these effects, with shifts in functional traits having implications for future ecosystem function.

The effects of phenoseason and storm characteristics on throughfall solute washoff and leaching dynamics from a temperate deciduous forest canopy.

Seasonal variations in the washoff and leaching dynamics of throughfall ionic fluxes represent a significant process affecting the biogeochemical cycling of forested ecosystems-particularly for temperate deciduous forests with distinct phenological seasons (or "phenoseasons"). Most studies on temperate deciduous forests aggregate seasonal throughfall fluxes to the leafed (growing) and leafless (dormant) periods, yet the phenological conditions controlling seasonality demand finer-scale demarcations that include the transitional phenoseasons (leaf senescence and emergence). To fill these gaps our study examines the washoff and leaching dynamics of Na(+), Mg(2+), K(+), Ca(2+), Cl(-), SO(4)(2-), and NO(3)(-) throughfall derived from bulk and sequentially sampled rain events across leafed, leafless and both transitional phenoseasons over a 3-year period (2008-2010). As throughfall washoff and leached solute fluxes are also closely-coupled to rainfall conditions, we further examine the effects of storm characteristics on phenoseasonal washoff-dominated (Na(+) and Cl(-)) and leaching-dominated (K(+), Ca(2+), Mg(2+)) fluxes through intrastorm event comparison plots and factorial MANOVA. Highly significant differences in leached and washoff solute fluxes were found across meteorological conditions (p<0.001) nested within phenoseasonal divisions (p<0.00001). Phenoseasonal washoff Na(+) and Cl(-) fluxes seemed to be more closely related to leaf area; whereas, leaching flux and canopy exchange of all solutes to correspond more with major phenological changes (when the canopies tend to be most metabolically active). The greatest differences in leached Mg(2+), K(+), Ca(2+), and SO(4)(2-) fluxes were not between the full leafed and leafless phenoseasons (33-80% difference), but between the transitional periods (80 to 200 fold greater during leaf senescence than leaf emergence). Intrastorm average canopy NO(3)(-) leaching, however, ranged from low losses (1 µmol(c)m(-2)h(-1)) to canopy uptake (-2 µmol(c)m(-2)h(-1)) during both transitional phenoseasons. K(+), Ca(2+), Mg(2+) were all markedly more exchangeable during senescence, with Ca(2+) and Mg(2+) being more tightly held by the canopy. Leaching rates and fluxes for all measured solutes were negligible to negative during emergence, except for K(+) and SO(4)(2-). Our results indicate that much of the variance in timing and magnitude of throughfall solute fluxes to forest soils within temperate deciduous ecosystems may be ascribed to phenologically-delineated seasons and storm conditions.