Respuesta de las comunidades microbianas a fluctuaciones de nutrientes en lagos de alta montaña

Resumen

Cold, oligotrophic lakes hold a large diversity of planktonic microbial organisms, protists and prokaryotes. The coexistence of such a large variety of organisms and living forms has challenged the ecological theory. The environmental fluctuations, particularly the contrast between seasons, have been pointed as an opportunity for coexistence. As the bases of this thesis, it was hypothesized that episodic nutrient enrichments play a major role in the maintenance of such high microbial diversity, particularly because nutrient loadings throughout the year consist in different amounts and combinations of phosphorus and nitrogen compounds, providing an opportunity for nutrient niche differentiation. Two approaches were used to investigate the planktonic microbial coexistence: field regular observations and experimental field manipulations. The first involved the monitoring of one annual cycle of the bacterioplankton community in a seasonally frozen lake and the second, an in situ nutrient enrichment experiment using enclosures. Both approaches were carried out in Lake Redon, an ultraoligotrophic deep high mountain lake that is ice-covered for about half a year. In Chapter 2, the seasonal changes of the bacterioplankton community are described. In Chapter 3 and 4, the protist- and bacteria-specific responses to episodic nutrient enrichments are investigated using self-filling enclosures and P and N additions. A new paradigm for bacterioplankton seasonal changes in seasonally ice-covered lakes is suggested in Chapter 2. In contrast to the traditionally-assumed assemblage seasonal replacement, there is a core of bacterioplankton assemblages developing fundamentally during the ice-covered period, which is resistant or resilient to the high irradiances conditions of the summer season. A different assemblage develops in the epilimnion during the ice-free season for a short period, taking the opportunity of unsuitable conditions for the core assemblages that proliferate in most of the water column during most of the year. The ENEX experiment aimed to investigate the relationship between episodic nutrient enrichments and species coexistence. Concerning the protist community (Chapter 3), P was the limiting nutrient promoting species competition, while N was involved in niche stabilising mechanisms. Most of the species typically growing at different periods of the year were recovered during the experiment from epilimnetic waters, showing that, although at undetectable abundances, species have the capacity to persist for long periods in the water column and quickly recover from low densities when favourable nutrient conditions are present. Concerning bacterioplankton (Chapter 4), there was also marked biomass increase to P enrichment. However, the comparison with the natural cycle in the lake showed that most of the response was mediated by the phytoplankton growth and the higher availability of labile organic carbon. There was little segregation at high taxonomic ranks (i.e., classes) of the bacteria across nutrient conditions; however, OTUs from the same family segregated across the P gradient and N form sources indicating niche differentiation that can promote coexistence. The thesis demonstrates the key role of episodic nutrient enrichments of varying composition for maintaining long-term stable non-equilibrium coexistence in the planktonic microbial community.