Background and aims Ancient Amazon soils are characterised by low concentrations of soil phosphorus (P). Therefore, it is hypothesised that plants may invest a substantial proportion of their resources belowground to adjust their P-uptake strategies, including root morphological, physiological (phosphatase enzyme activities) and biotic (arbuscular mycorrhizal (AM) associations) adaptations. Since these strategies are energy demanding, we hypothesise that trade-offs between morphological traits and root phosphatase exudation and symbiotic associations would occur. Specifically, we expected that plants which invest in finer roots, and therefore have greater ability to explore large soil volumes, would have a high investment in physiological adaptations such as enhanced phosphatase production. In contrast, we expected that plants with predominantly thicker roots would invest more in symbiotic associations, in which carbon is traded for P acquired from AM fungal communities. Methods We collected absorptive roots (<2 mm diameter) from a lowland Central Amazon forest near Manaus, Brazil. We measured fine root diameter, specific root length (SRL), specific root area (SRA), root tissue density (RTD), root phosphatase activity (APase) and arbuscular mycorrhizal (AM) fungi colonisation. Results Root morphological traits were related to APase activity, with higher APase activity in roots with higher SRL and SRA but lower RTD. However, the degree of AM colonisation was not related to any measured root morphological trait. Conclusions Fine absorptive roots likely benefit from having low RTD, high SRL, SRA and APase exudation to acquire P efficiently. However, because AM colonisation was not related to root morphology, we suggest that investment in multiple P-uptake strategies is required for maintaining productivity in Central Amazon forests.
Summary
Soil nutrient availability can strongly affect root traits. In tropical forests, phosphorus (P) is often considered the main limiting nutrient for plants. However, support for the P paradigm is limited, and N and cations might also control tropical forests functioning.
We used a large‐scale experiment to determine how the factorial addition of nitrogen (N), P and cations affected root productivity and traits related to nutrient acquisition strategies (morphological traits, phosphatase activity, arbuscular mycorrhizal colonisation and nutrient contents) in a primary rainforest growing on low‐fertility soils in Central Amazonia after 1 yr of fertilisation.
Multiple root traits and productivity were affected. Phosphorus additions increased annual root productivity and root diameter, but decreased root phosphatase activity. Cation additions increased root productivity at certain times of year, also increasing root diameter and mycorrhizal colonisation. P and cation additions increased their element concentrations in root tissues. No responses were detected with N addition.
Here we showed that rock‐derived nutrients determined root functioning in low‐fertility Amazonian soils, demonstrating not only the hypothesised importance of P, but also highlighting the role of cations. The changes in fine root traits and productivity indicated that even slow‐growing tropical rainforests can respond rapidly to changes in resource availability.
The productivity of rainforests growing on highly-weathered tropical soils is expected to be limited by phosphorus (P) availability 1 . Yet, controlled fertilisation experiments have failed to demonstrate a dominant role for P in controlling tropical forest net primary productivity (NPP). Recent syntheses have demonstrated that responses to N addition are as large as to P 2 , and adaptations to low P availability appear to allow NPP to be maintained across major soil P gradients 3 . Thus, the extent to which P availability limits tropical forest productivity is highly uncertain. The majority of the Amazonia, however, is characterised by soils even more depleted in P than where most tropical fertilisation experiments have previously taken place 2 . Thus, we established the first P, nitrogen (N), and base cation addition experiment in an old growth Amazon rainforest, with the site's low soil P content representative of ~60% of the basin. Here we show that NPP increased exclusively with P addition. After 2 years, strong responses were observed in fine root (+29%) and canopy productivity (+19%), but not stem growth. The direct evidence of P limitation of NPP suggests that P availability may restrict Amazon forest responses to CO2 fertilisation 4 , with major implications for future carbon sequestration and forest resilience to climate change.
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