Tree growth and management in Ugandan agroforestry systems: effects of root pruning on tree growth and crop yield

Tree root pruning is a potential tool for managing below-ground competition when trees and crops are grown together in agroforestry systems. This study investigates its effects on growth and root distribution of Alnus acuminata (HB & K), Casuarina equisetifolia (L), Grevillea robusta (A. Cunn. ex R. Br), Maesopsis eminii (Engl.), and Markhamia lutea (Benth.) K. Schum. and on yield of adjacent crops in sub-humid Uganda. The trees were 3 years old at the commencement of the study, and most species were competing strongly with crops. Tree roots were pruned 41 months after planting by cutting and back-filling a trench to a depth of 0.3 m, at a distance of 0.3 m from the trees, on one side of the tree row. The trench was re-opened and roots re-cut at 50 and 62 months after planting. Effects on tree growth and root distribution were assessed over a 3 year period, and crop yield after the third root pruning at 62 months is reported here. Overall, root pruning had only a slight effect on tree growth: height growth was unaffected and diameter growth was reduced by only 4 %. A substantial amount of root re-growth was observed by 11 months after pruning. Tree species varied in the number and distribution of their roots, and Casuarina and Markhamia had considerably more roots per unit of trunk volume than the other tree species, especially in the surface soil layers. Casuarina and Maesopsis were the most competitive tree species with crops and Grevillea and Markhamia the least. Crop yield data provides strong evidence of the redistribution of root activity following root pruning, so that competition increased on the unpruned side of tree rows. Thus, one-sided root pruning will only be of use to farmers in a few circumstances. Key words: Alnus acuminata, Casuarina equisetifolia, Grevillea robusta, Maesopsis eminii, Markhamia lutea, root distribution, root functio

Legumes are different:Leaf nitrogen, photosynthesis, and water use efficiency

Using robust, pairwise comparisons and a global dataset, we show that nitrogen concentration per unit leaf mass for nitrogen-fixing plants (N(2)FP; mainly legumes plus some actinorhizal species) in nonagricultural ecosystems is universally greater (43–100%) than that for other plants (OP). This difference is maintained across Koppen climate zones and growth forms and strongest in the wet tropics and within deciduous angiosperms. N(2)FP mostly show a similar advantage over OP in nitrogen per leaf area (N(area)), even in arid climates, despite diazotrophy being sensitive to drought. We also show that, for most N(2)FP, carbon fixation by photosynthesis (A(sat)) and stomatal conductance (g(s)) are not related to N(area)—in distinct challenge to current theories that place the leaf nitrogen–A(sat) relationship at the center of explanations of plant fitness and competitive ability. Among N(2)FP, only forbs displayed an N(area)–g(s) relationship similar to that for OP, whereas intrinsic water use efficiency (WUE(i); A(sat)/g(s)) was positively related to N(area) for woody N(2)FP. Enhanced foliar nitrogen (relative to OP) contributes strongly to other evolutionarily advantageous attributes of legumes, such as seed nitrogen and herbivore defense. These alternate explanations of clear differences in leaf N between N(2)FP and OP have significant implications (e.g., for global models of carbon fluxes based on relationships between leaf N and A(sat)). Combined, greater WUE and leaf nitrogen—in a variety of forms—enhance fitness and survival of genomes of N(2)FP, particularly in arid and semiarid climates

Legume abundance along successional and rainfall gradients in Neotropical forests

The nutrient demands of regrowing tropical forests are partly satisfied by nitrogen-fixing legume trees, but our understanding of the abundance of those species is biased towards wet tropical regions. Here we show how the abundance of Leguminosae is affected by both recovery from disturbance and large-scale rainfall gradients through a synthesis of forest inventory plots from a network of 42 Neotropical forest chronosequences. During the first three decades of natural forest regeneration, legume basal area is twice as high in dry compared with wet secondary forests. The tremendous ecological success of legumes in recently disturbed, water-limited forests is likely to be related to both their reduced leaflet size and ability to fix N2, which together enhance legume drought tolerance and water-use efficiency. Earth system models should incorporate these large-scale successional and climatic patterns of legume dominance to provide more accurate estimates of the maximum potential for natural nitrogen fixation across tropical forests.Additional co-authors: Rebecca J. Cole, Gabriel Dalla Colletta, Ben de Jong, Julie S. Denslow, Saara J. DeWalt, Juan Manuel Dupuy, Sandra M. Durán, Mário Marcos do Espírito Santo, G. Wilson Fernandes, Yule Roberta Ferreira Nunes, Bryan Finegan, Vanessa Granda Moser, Jefferson S. Hall, José Luis Hernández-Stefanoni, André B. Junqueira, Deborah Kennard, Edwin Lebrija-Trejos, Susan G. Letcher, Madelon Lohbeck, Erika Marín-Spiotta, Miguel Martínez-Ramos, Jorge A. Meave, Duncan N. L. Menge, Francisco Mora, Rodrigo Muñoz, Robert Muscarella, Susana Ochoa-Gaona, Edith Orihuela-Belmonte, Rebecca Ostertag, Marielos Peña-Claros, Eduardo A. Pérez-García, Daniel Piotto, Peter B. Reich, Casandra Reyes-García, Jorge Rodríguez-Velázquez, I. Eunice Romero-Pérez, Lucía Sanaphre-Villanueva, Arturo Sanchez-Azofeifa, Naomi B. Schwartz, Arlete Silva de Almeida, Jarcilene S. Almeida-Cortez, Whendee Silver, Vanessa de Souza Moreno, Benjamin W. Sullivan, Nathan G. Swenson, Maria Uriarte, Michiel van Breugel, Hans van der Wal, Maria das Dores Magalhães Veloso, Hans F. M. Vester, Ima Célia Guimarães Vieira, Jess K. Zimmerman & Jennifer S. Power

The innovation of the symbiosome has enhanced the evolutionary stability of nitrogen fixation in legumes

Nitrogen-fixing symbiosis is globally important in ecosystem functioning and agriculture, yet the evolutionary history of nodulation remains the focus of considerable debate. Recent evidence suggesting a single origin of nodulation followed by massive parallel evolutionary losses raises questions about why a few lineages in the N2 -fixing clade retained nodulation and diversified as stable nodulators, while most did not. Within legumes, nodulation is restricted to the two most diverse subfamilies, Papilionoideae and Caesalpinioideae, which show stable retention of nodulation across their core clades. We characterize two nodule anatomy types across 128 species in 56 of the 152 genera of the legume subfamily Caesalpinioideae: fixation thread nodules (FTs), where nitrogen-fixing bacteroids are retained within the apoplast in modified infection threads, and symbiosomes, where rhizobia are symplastically internalized in the host cell cytoplasm within membrane-bound symbiosomes (SYMs). Using a robust phylogenomic tree based on 997 genes from 147 Caesalpinioideae genera, we show that losses of nodulation are more prevalent in lineages with FTs than those with SYMs. We propose that evolution of the symbiosome allows for a more intimate and enduring symbiosis through tighter compartmentalization of their rhizobial microsymbionts, resulting in greater evolutionary stability of nodulation across this species-rich pantropical legume clade