<p>Nutrient availability in tropical forest ecosystems plays a critical role in sustaining forest growth and productivity. Observational evidence for nutrient limitations on net primary productivity (NPP) in the tropics is rare yet crucial for predicting the impacts of human-induced changes on tropical forests, particularly for underrepresented tropical regions in Africa. In an ecosystem-scale nutrient manipulation experiment, we assessed the response of different components of above-ground net primary production (ANPP) to nutrient addition of nitrogen (N), phosphorus (P), potassium (K) and all possible combinations (NP, NK, PK, and NPK) at rates of 125 kg N ha<sup>-1</sup>yr<sup>-1</sup>, 50 kg P ha<sup>-1</sup> yr<sup>-1</sup> and 50 kg K ha<sup>-1</sup>yr<sup>-1</sup>.</p><p>We established 32 (8 treatments &#215; 4 replicates) experimental plots of 40 &#215; 40 m<sup>2</sup> each and measured stem growth of over 15,000 trees with diameter at breast height (dbh) &#8805; 1 cm as well as litter production and above-ground woody biomass production (AWBP), of a lower-montane tropical forest (1100 m a.s.l.) in northwestern Uganda.</p><p>After 18 months of nutrient addition, we found that different aspects of ANPP, including litter production and AWBP are controlled by multiple soil nutrients. Specifically, we measured higher total fine-litter production in the N (13.6 &#177; 1.4 Mg ha<sup>-1 </sup>yr<sup>-1</sup>) and K (13.3 &#177; 1.8 Mg ha<sup>-1 </sup>yr<sup>-1</sup>) addition plots than the control (11.1 &#177; 0.6 Mg ha<sup>-1 </sup>yr<sup>-1</sup>) plots. Both reproductive litter (flowers and fruits; 10% of total fine-litter fall) and leaf litter (62% of total fine-litter fall) significantly increased with K addition. In general, fine-litter production in our plots is higher than what has been reported so far for lower-montane tropical forests. Increased AWBP is associated with N addition plots. The response of trees to nutrient addition however, varied with tree sizes. Trees with dbh between 10 &#8211; 30 cm increased significantly in AWBP under PK addition. There was no effect of nutrient addition associated with either smaller (1 &#8211; 10 cm dbh) or larger trees (dbh > 30 cm). The medium-sized trees which may have experienced resource competition but have now transitioned into the canopy layer (exposed to sunlight) are able to use additional nutrient for active growth. In contrast, bigger trees may allocate extra nutrient for reproduction and leaf-vitality, while smaller trees remain shaded, co-limited by sunlight and therefore unable to utilize increased available nutrients for stem diameter growth. ANPP increased by 39% with N addition and marginally by 23% with K additions relative to the control. In conclusion, our experiment provides evidence of N and potentially K limitation of ANPP in this lower-montane tropical forest, and highlights that, in a highly diverse ecosystem different components of ANPP may be regulated by multiple nutrients.&#160;</p>
Plant-soil associations in a lower montane tropical forest: physiological acclimation and herbivore-mediated responses to nitrogen addition