Abstract
Root biomass distributions have long been used to infer patterns of resource uptake. These patterns are used to understand plant growth, plant coexistence and water budgets. Root biomass, however, may be a poor indicator of resource uptake because large roots typically do not absorb water, fine roots do not absorb water from dry soils and roots of different species can be difficult to differentiate. In a sub-tropical savanna, Kruger Park, South Africa, we used a hydrologic tracer experiment to describe the abundance of active grass and tree roots across the soil profile. We then used this tracer data to parameterize a water movement model (Hydrus 1D). The model accounted for water availability and estimated grass and tree water uptake by depth over a growing season. Most root biomass was found in shallow soils (0–20 cm) and tracer data revealed that, within these shallow depths, half of active grass roots were in the top 12 cm while half of active tree roots were in the top 21 cm. However, because shallow soils provided roots with less water than deep soils (20–90 cm), the water movement model indicated that grass and tree water uptake was twice as deep as would be predicted from root biomass or tracer data alone: half of grass and tree water uptake occurred in the top 23 and 43 cm, respectively. Niche partitioning was also greater when estimated from water uptake rather than tracer uptake. Contrary to long-standing assumptions, shallow grass root distributions absorbed 32% less water than slightly deeper tree root distributions when grasses and trees were assumed to have equal water demands. Quantifying water uptake revealed deeper soil water uptake, greater niche partitioning and greater benefits of deep roots than would be estimated from root biomass or tracer uptake data alone.
Highlights
Plant uptake of soil resources is one of the most fundamental processes of life on earth
In arid and semi-arid systems, which represent nearly half of terrestrial ecosystems, plant growth is highly sensitive to soil water availability [4, 5] and plant coexistence is believed to reflect differences in PLOS ONE | DOI:10.1371/journal.pone
Walter’s two-layer hypothesis suggests that grasses predominate in savannas due to greater water use efficiency and dense shallow roots but that woody plants can coexist with grasses due to deep roots [1, 8]. This hypothesis was developed for drier savannas (i.e., < 500 mm precipitation), but it is often applied to a wide range of arid and semi-arid ecosystems [8]. Testing this hypothesis requires measurements of the location, timing and extent of water uptake by grasses and trees in the field, yet much of the support for the two-layer hypothesis continues to be inferred from observations of root distributions and not plant water uptake [6, 8,9,10]
Summary
Plant uptake of soil resources is one of the most fundamental processes of life on earth. Our general approach was to 1) inject the hydrologic tracer deuterium oxide (D2O) into five soil depths, three times during a growing season in a sub-tropical savanna to determine the proportion of tracer uptake from each depth by dominant grasses and trees [12, 27], 2) parameterize Hydrus 1D, a widely-used soil water movement model [15], with tracer-derived estimates of active root distributions. This approach produced modeled estimates of the amount of water removed by grasses and trees from different soil depths over a growing season [22, 29]. Broadly, critical components of this modeling approach, namely aerodynamic resistances and preferential flow paths in the soil will need to be validated
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