Phosphorus Limitation During Long-Term Ecosystem Development
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Phosphorus limitation during long-term ecosystem development Benjamin L. Turner1, Andrew Jones2, Etienne Laliberté3, Ed Vicenzi4, Jeremy Drake5 1Smithsonian Tropical Research Institute, Republic of Panama 2Oregon State University, Corvallis, OR, USA 3University of Western Australia, Crawley, Australia 4Smithsonian Museum Conservation Institute, Suitland, MD, USA 5Smithsonian Astrophysical Observatory, Cambridge, MA, USA The final frontier? “We know more about the movement of celestial bodies than about the soil underfoot.” Leonardo da Vinci, c. 1500 Soils and terrestrial ecology • Soils support terrestrial life – soils provide the structural support for plants – soils regulate water supply – soils provide a reservoir of nutrients • Soils are biologically diverse – a handful of soil contains tens of thousands of distinct microbial species • How do soils influence the productivity, diversity, and distribution of organisms in the environment? Phosphorus: The Devil’s Element! • Fundamental to life on earth – protein synthesis (RNA) – passage of genetic information (DNA) – cell membranes (phospholipids) – energy transfer (ATP) • Limitation of plant productivity – widespread phosphorus deficiency in both terrestrial and aquatic ecosystems • Peak phosphorus? – essential in modern agriculture, but are phosphorus reserves running out? Climate Organisms Relief Parent material Soil forming factors forming Soil Aquic Hapludult, Time SERC, Maryland Increasing soil age Soil chronosequence in coastal dunes at Haast, New Zealand 300 years old 4000 years old Changes in nutrient status during ecosystem development N-limitation P-limitation Nitrogen Nutrient content Nutrient Phosphorus Time Phosphorus transformations during pedogenesis (Walker and Syers, 1976) Old soils contain only occluded and organic phosphorus Total P Ca-P “Non-occluded” P Phosphorus content Phosphorus “Occluded” P Organic P Time 1.0 Total Phosphorus (g kg-1) 0.5 0.0 Total Nitrogen (g kg-1) 10 5 0 40 N:P Ratio 20 0 The Franz Josef Glacier, New Zealand 100 101 102 103 104 105 106 Years Franz Josef Glacier Arawhata, New Zealand Cooloola, Australia Northern Arizona Volcanic Field Jurien Bay, Western Australia Hawaiian Islands Karangarua Terraces, New Zealand Mendocino Staircase, California Haast Dunes, New Zealand Foliar phosphorus along the Jurien Bay soil chronosequence, Western Australia ) 1 - (µg P g (µg P Foliarphosphorus 102 103 104 105 106 Approximate soil age (years) Young Old chronosequence, Australia eastern Cooloola Shift towards stress-tolerant tree species along the Franz Josef chronosequence Angiosperms (e.g.): Angiosperms Weinmannia racemosa Podocarpaceae Dicksonia squarrosa 100 Podocarpaceae (e.g.): Dacrydium cupressinum Podocarpus hallii 75 50 25 Canopy (%) Cover 0 100 101 102 103 104 105 Years Plant strategies for acquiring soil phosphorus • Synthesis of phosphatase enzymes – a ubiquitous response of plants to the need for phosphorus • Formation of mycorrhizal symbioses – some are extremely efficient at acquiring soil phosphorus • Cluster roots and organic anions – compounds like citrate can solubilize large amounts of soil phosphorus Changes in plant community composition during ecosystem development From: Lambers et al. (2008) Trends in Ecology and Evolution 23, 95–103. Greater plant diversity on ancient landscapes 102 103 104 105 106 Approximate soil age (years) Significance of the soil microbial biomass • Large phosphorus pool in microbial biomass • Microbial phosphorus > plant phosphorus for most of the sequence • Intense plant–microbe competition for phosphorus on old soils? Changes in microbial communities with soil age 1.0 Mineral • Increase in fungal to bacterial ratios Organic 0.8 – e.g., in mineral and organic horizons along the Franz Josef chronosequence 0.6 0.4 Fungal:bacterial ratio 0.2 • Investment in phosphorus acquisition Fungal to bacterial ratio bacterialto Fungal 0.0 – e.g., for two phosphatase enzymes along 3 4 5 6 7 8 9 10 11 12 13 Chronosequenceln (yr) stage the Haast chronosequence 2.0 25 Mineral Organic 1.8 Phosphomonoesterase ) Phosphodiesterase (x5) 1 - 20 1.6 min 1 activity • Change in microbial communities - 151.4 1.2 – decline in bacterial diversity and richness 10 along the New Zealand chronosequences g product 1.0 Gram +ve:Gram -ve ratio 5 0.8 nmol ( Phosphatase 0.60 3 14 25 36 4 7 5 8 6 9 7 108 119 1210 13 ln (yr) Bacterial community composition during pedogenesis • Relative abundance of bacterial phyla along the Franz Josef chronosequence Actinomycetes α -Proteobacteria Bacilli 50 40 4 40 30 30 20 2 20 10 10 0 0 0 β-Proteobacteria γ-Proteobacteria Bacteroidetes 12 6 8 10 r=0.88 6 8 4 Relative abundance (%) abundance Relative r=0.85 6 4 4 2 2 2 0 0 0 101 102 103 104 105 101 102 103 104 105 101 102 103 104 105 Soil age (years) Ancient soils: an extreme environment for microbes? Summary: relevance to astrobiology? • Long-term decline in phosphorus availability – extremely low phosphorus concentrations in old soils – phosphorus rejuvenation by tectonic activity (or other major disturbance) • Consequences for terrestrial life – intense biological phosphorus limitation – decline in productivity, greater specialization, greater diversity (plants) • Life on ancient terrestrial landscapes as an analogue for low- tectonic worlds?.