Joint environmental and social benefits from diversified agriculture.

Agricultural simplification continues to expand at the expense of more diverse forms of agriculture. This simplification, for example, in the form of intensively managed monocultures, poses a risk to keeping the world within safe and just Earth system boundaries. Here, we estimated how agricultural diversification simultaneously affects social and environmental outcomes. Drawing from 24 studies in 11 countries across 2655 farms, we show how five diversification strategies focusing on livestock, crops, soils, noncrop plantings, and water conservation benefit social (e.g., human well-being, yields, and food security) and environmental (e.g., biodiversity, ecosystem services, and reduced environmental externalities) outcomes. We found that applying multiple diversification strategies creates more positive outcomes than individual management strategies alone. To realize these benefits, well-designed policies are needed to incentivize the adoption of multiple diversification strategies in unison.

Water status dynamics and drought tolerance of juvenile European beech, Douglas fir and Norway spruce trees as dependent on neighborhood and nitrogen supply.

To increase the resilience of forests to drought and other hazards, foresters are increasingly planting mixed stands. This requires knowledge about the drought response of tree species in pure- and mixed-culture neighborhoods. In addition, drought frequently interacts with continued atmospheric nitrogen (N) deposition. To disentangle these factors for European beech, Norway spruce and Douglas fir, we conducted a replicated three-factorial sapling growth experiment with three moisture levels, (high, medium and low), two N levels (high and ambient) and pure and mixed-culture neighborhoods. We measured biomass, stomatal conductance (GS), shoot water potential (at predawn: ΨPD, midday, and turgor loss point: ΨTLP), branch xylem embolism resistance (Ψ50), and minimum epidermal conductance (Gmin). The three species differed most with respect to Gmin (10-fold higher in beech than in the conifers), hydroscape area (larger in beech), and the time elapsed to reach stomatal closure (TΨGS90) and ΨTLP (TTLP; shorter in beech), while Ψ50 and ΨTLP were remarkably similar. Neighborhood (pure vs. mixed-culture) influenced biomass production, water status and hydraulic traits, notably GS (higher in Douglas fir, but lower in spruce and beech, in mixtures than pure culture), hydraulic safety margin (smaller for beech in mixtures), and TΨGS90 and TTLP (shorter for spruce in mixture). High N generally increased GS, but no consistent N effects on leaf water status and hydraulic traits were detected, suggesting that neighbor identity had a larger effect on plant water relations than N availability. We conclude that both tree neighborhood and N availability modulate the drought response of beech, spruce and Douglas fir. Species mixing can alleviate the drought stress of some species, but often by disadvantaging other species. Thus, our study suggests stabilizing and building resilience of production forests against a drier and warmer climate may depend primarily on the right species choice; species mixing can support the agenda.

Balancing economic and ecological functions in smallholder and industrial oil palm plantations.

The expansion of the oil palm industry in Indonesia has improved livelihoods in rural communities, but comes at the cost of biodiversity and ecosystem degradation. Here, we investigated ways to balance ecological and economic outcomes of oil palm cultivation. We compared a wide range of production systems, including smallholder plantations, industrialized company estates, estates with improved agronomic management, and estates with native tree enrichment. Across all management types, we assessed multiple indicators of biodiversity, ecosystem functions, management, and landscape structure to identify factors that facilitate economic-ecological win-wins, using palm yields as measure of economic performance. Although, we found that yields in industrialized estates were, on average, twice as high as those in smallholder plantations, ecological indicators displayed substantial variability across systems, regardless of yield variations, highlighting potential for economic-ecological win-wins. Reducing management intensity (e.g., mechanical weeding instead of herbicide application) did not lower yields but improved ecological outcomes at moderate costs, making it a potential measure for balancing economic and ecological demands. Additionally, maintaining forest cover in the landscape generally enhanced local biodiversity and ecosystem functioning within plantations. Enriching plantations with native trees is also a promising strategy to increase ecological value without reducing productivity. Overall, we recommend closing yield gaps in smallholder cultivation through careful intensification, whereas conventional plantations could reduce management intensity without sacrificing yield. Our study highlights various pathways to reconcile the economics and ecology of palm oil production and identifies management practices for a more sustainable future of oil palm cultivation.

Ambient and substrate energy influence decomposer diversity differentially across trophic levels.

The species-energy hypothesis predicts increasing biodiversity with increasing energy in ecosystems. Proxies for energy availability are often grouped into ambient energy (i.e., solar radiation) and substrate energy (i.e., non-structural carbohydrates or nutritional content). The relative importance of substrate energy is thought to decrease with increasing trophic level from primary consumers to predators, with reciprocal effects of ambient energy. Yet, empirical tests are lacking. We compiled data on 332,557 deadwood-inhabiting beetles of 901 species reared from wood of 49 tree species across Europe. Using host-phylogeny-controlled models, we show that the relative importance of substrate energy versus ambient energy decreases with increasing trophic levels: the diversity of zoophagous and mycetophagous beetles was determined by ambient energy, while non-structural carbohydrate content in woody tissues determined that of xylophagous beetles. Our study thus overall supports the species-energy hypothesis and specifies that the relative importance of ambient temperature increases with increasing trophic level with opposite effects for substrate energy.

A meta-analysis of responses in floral traits and flower-visitor interactions to water deficit.

Alterations in water availability and drought events as predicted by climate change scenarios will increasingly impact natural communities with effects already emerging at present. Water deficit leads to increasing physiological stress in plants, likely affecting floral development and causing changes in floral morphology, nectar and pollen production or scent. Understanding how these floral traits are altered by water deficit is necessary to predict changes in plant-pollinator interactions and how communities are impacted in the future. Here we employ a meta-analysis approach to synthesize the current evidence of experimental water deficit on floral traits and plant-pollinator interactions. Furthermore, we explore experimental factors potentially increasing heterogeneity between studies and provide ideas how to enhance comparability between studies. In the end, we highlight future directions and knowledge gaps for floral traits and plant-pollinator interactions under water deficit. Our analysis showed consistent decreases in floral size, number of flowers and nectar volume to reduced water availability. Other floral traits such as the start of flowering or herkogamy showed no consistent pattern. This indicates that effects of reduced water availability differ between specific traits that are potentially involved in different functions such as pollinator attraction or efficiency. We found no general decreasing visitation rates with water deficit for flower-visitor interactions. Furthermore, the comparison of available studies suggests that increased reporting of plant stress severity and including more hydraulic and physiological measurements will improve the comparability across experiments and aid a more mechanistic understanding of plant-pollinator interactions under altered environmental conditions. Overall, our results show that water deficit has the potential to strongly affect plant-pollinator interactions via changes in specific floral traits. Linking these changes to pollination services and pollinator performance is one crucial step for understanding how changing water availability and drought events under climate change will alter plant and pollinator communities.

Trade-offs between multifunctionality and profit in tropical smallholder landscapes.

Land-use transitions can enhance the livelihoods of smallholder farmers but potential economic-ecological trade-offs remain poorly understood. Here, we present an interdisciplinary study of the environmental, social and economic consequences of land-use transitions in a tropical smallholder landscape on Sumatra, Indonesia. We find widespread biodiversity-profit trade-offs resulting from land-use transitions from forest and agroforestry systems to rubber and oil palm monocultures, for 26,894 aboveground and belowground species and whole-ecosystem multidiversity. Despite variation between ecosystem functions, profit gains come at the expense of ecosystem multifunctionality, indicating far-reaching ecosystem deterioration. We identify landscape compositions that can mitigate trade-offs under optimal land-use allocation but also show that intensive monocultures always lead to higher profits. These findings suggest that, to reduce losses in biodiversity and ecosystem functioning, changes in economic incentive structures through well-designed policies are urgently needed.

Parenchyma Abundance in Wood of Evergreen Trees Varies Independently of Nutrients.

The abundance of living cells in wood-mainly as interconnected axial and ray parenchyma networks-varies widely between species. However, the functional significance of this variation and its role in plant ecological strategies is poorly understood, as is the extent to which different parenchyma fractions are favored in relation to soil nutrients and hydraulic functions. We analyzed wood tissue fractions of 16 Australian angiosperm species sampled from two nearby areas with similar climate but very different soil nutrient profiles and investigated structure-function links with soil and tissue nutrient concentrations and other plant traits. We expected the variation in parenchyma fractions to influence nutrient concentrations in wood xylem, and to find species with lower parenchyma fractions and accordingly lower nutrient requirements on lower-nutrient soils. Surprisingly, both axial and ray parenchyma fractions were mostly unrelated to tissue and soil nutrient concentrations, except for nitrogen concentration in stem sapwood. Species from low nutrient soils showed higher fractional P translocation from both leaves and sapwood, but little patterning with respect to tissue nitrogen. While species from high and low nutrient soils clearly clustered along the soil-fertility axis, their tissue composition varied independently from plant functional traits related to construction costs and hydraulic anatomy. Our findings imply that there is considerable variation among species in the nutrient concentrations within different parenchyma tissues. The anatomical composition of wood tissue seems unrelated to plant nutrient requirements. Even though xylem parenchyma is involved in metabolic functions such as nutrient translocation and storage, parenchyma abundance on its own does not directly explain variation in these functions, even in co-occurring species. While parenchyma is highly abundant in wood of angiosperm trees, we are still lacking a convincing ecological interpretation of its variability and role in whole-tree nutrient budgets.

High porosity with tiny pore constrictions and unbending pathways characterize the 3D structure of intervessel pit membranes in angiosperm xylem.

Pit membranes between xylem vessels play a major role in angiosperm water transport. Yet, their three-dimensional (3D) structure as fibrous porous media remains unknown, largely due to technical challenges and sample preparation artefacts. Here, we applied a modelling approach based on thickness measurements of fresh and fully shrunken pit membranes of seven species. Pore constrictions were also investigated visually by perfusing fresh material with colloidal gold particles of known sizes. Based on a shrinkage model, fresh pit membranes showed tiny pore constrictions of ca. 20 nm, but a very high porosity (i.e. pore volume fraction) of on average 0.81. Perfusion experiments showed similar pore constrictions in fresh samples, well below 50 nm based on transmission electron microscopy. Drying caused a 50% shrinkage of pit membranes, resulting in much smaller pore constrictions. These findings suggest that pit membranes represent a mesoporous medium, with the pore space characterized by multiple constrictions. Constrictions are much smaller than previously assumed, but the pore volume is large and highly interconnected. Pores do not form highly tortuous, bent, or zigzagging pathways. These insights provide a novel view on pit membranes, which is essential to develop a mechanistic, 3D understanding of air-seeding through this porous medium.

Is xylem of angiosperm leaves less resistant to embolism than branches? Insights from microCT, hydraulics, and anatomy.

According to the hydraulic vulnerability segmentation hypothesis, leaves are more vulnerable to decline of hydraulic conductivity than branches, but whether stem xylem is more embolism resistant than leaves remains unclear. Drought-induced embolism resistance of leaf xylem was investigated based on X-ray microcomputed tomography (microCT) for Betula pendula, Laurus nobilis, and Liriodendron tulipifera, excluding outside-xylem, and compared with hydraulic vulnerability curves for branch xylem. Moreover, bordered pit characters related to embolism resistance were investigated for both organs. Theoretical P50 values (i.e. the xylem pressure corresponding to 50% loss of hydraulic conductance) of leaves were generally within the same range as hydraulic P50 values of branches. P50 values of leaves were similar to branches for L. tulipifera (-2.01 versus -2.10 MPa, respectively), more negative for B. pendula (-2.87 versus -1.80 MPa), and less negative for L. nobilis (-6.4 versus -9.2 MPa). Despite more narrow conduits in leaves than branches, mean interconduit pit membrane thickness was similar in both organs, but significantly higher in leaves of B. pendula than in branches. This case study indicates that xylem shows a largely similar embolism resistance across leaves and branches, although differences both within and across organs may occur, suggesting interspecific variation with regard to the hydraulic vulnerability segmentation hypothesis.

Carbon costs and benefits of Indonesian rainforest conversion to plantations.

Land-use intensification in the tropics plays an important role in meeting global demand for agricultural commodities but generates high environmental costs. Here, we synthesize the impacts of rainforest conversion to tree plantations of increasing management intensity on carbon stocks and dynamics. Rainforests in Sumatra converted to jungle rubber, rubber, and oil palm monocultures lost 116 Mg C ha-1, 159 Mg C ha-1, and 174 Mg C ha-1, respectively. Up to 21% of these carbon losses originated from belowground pools, where soil organic matter still decreases a decade after conversion. Oil palm cultivation leads to the highest carbon losses but it is the most efficient land use, providing the lowest ratio between ecosystem carbon storage loss or net primary production (NPP) decrease and yield. The imbalanced sharing of NPP between short-term human needs and maintenance of long-term ecosystem functions could compromise the ability of plantations to provide ecosystem services regulating climate, soil fertility, water, and nutrient cycles.

Vessel-associated cells in angiosperm xylem: Highly specialized living cells at the symplast-apoplast boundary.

BACKGROUND: Vessel-associated cells (VACs) are highly specialized, living parenchyma cells that are in direct contact with water-conducting, dead vessels. The contact may be sparse or in large tight groups of parenchyma that completely surrounds vessels. VACs differ from vessel distant parenchyma in physiology, anatomy, and function and have half-bordered pits at the vessel-parenchyma juncture. The distinct anatomy of VACs is related to the exchange of substances to and from the water-transport system, with the cells long thought to be involved in water transport in woody angiosperms, but where direct experimental evidence is lacking. SCOPE: This review focuses on our current knowledge of VACs regarding anatomy and function, including hydraulic capacitance, storage of nonstructural carbohydrates, symplastic and apoplastic interactions, defense against pathogens and frost, osmoregulation, and the novel hypothesis of surfactant production. Based on microscopy, we visually represent how VACs vary in dimensions and general appearance between species, with special attention to the protoplast, amorphous layer, and the vessel-parenchyma pit membrane. CONCLUSIONS: An understanding of the relationship between VACs and vessels is crucial to tackling questions related to how water is transported over long distances in xylem, as well as defense against pathogens. New avenues of research show how parenchyma-vessel contact is related to vessel diameter and a new hypothesis may explain how surfactants arising from VAC can allow water to travel under negative pressure. We also reinforce the message of connectivity between VAC and other cells between xylem and phloem.

Direct and cascading impacts of tropical land-use change on multi-trophic biodiversity.

The conversion of tropical rainforest to agricultural systems such as oil palm alters biodiversity across a large range of interacting taxa and trophic levels. Yet, it remains unclear how direct and cascading effects of land-use change simultaneously drive ecological shifts. Combining data from a multi-taxon research initiative in Sumatra, Indonesia, we show that direct and cascading land-use effects alter biomass and species richness of taxa across trophic levels ranging from microorganisms to birds. Tropical land use resulted in increases in biomass and species richness via bottom-up cascading effects, but reductions via direct effects. When considering direct and cascading effects together, land use was found to reduce biomass and species richness, with increasing magnitude at higher trophic levels. Our analyses disentangle the multifaceted effects of land-use change on tropical ecosystems, revealing that biotic interactions on broad taxonomic scales influence the ecological outcome of anthropogenic perturbations to natural ecosystems.

Land-use choices follow profitability at the expense of ecological functions in Indonesian smallholder landscapes.

Smallholder-dominated agricultural mosaic landscapes are highlighted as model production systems that deliver both economic and ecological goods in tropical agricultural landscapes, but trade-offs underlying current land-use dynamics are poorly known. Here, using the most comprehensive quantification of land-use change and associated bundles of ecosystem functions, services and economic benefits to date, we show that Indonesian smallholders predominantly choose farm portfolios with high economic productivity but low ecological value. The more profitable oil palm and rubber monocultures replace forests and agroforests critical for maintaining above- and below-ground ecological functions and the diversity of most taxa. Between the monocultures, the higher economic performance of oil palm over rubber comes with the reliance on fertilizer inputs and with increased nutrient leaching losses. Strategies to achieve an ecological-economic balance and a sustainable management of tropical smallholder landscapes must be prioritized to avoid further environmental degradation.

Ecological and socio-economic functions across tropical land use systems after rainforest conversion.

Tropical lowland rainforests are increasingly threatened by the expansion of agriculture and the extraction of natural resources. In Jambi Province, Indonesia, the interdisciplinary EFForTS project focuses on the ecological and socio-economic dimensions of rainforest conversion to jungle rubber agroforests and monoculture plantations of rubber and oil palm. Our data confirm that rainforest transformation and land use intensification lead to substantial losses in biodiversity and related ecosystem functions, such as decreased above- and below-ground carbon stocks. Owing to rapid step-wise transformation from forests to agroforests to monoculture plantations and renewal of each plantation type every few decades, the converted land use systems are continuously dynamic, thus hampering the adaptation of animal and plant communities. On the other hand, agricultural rainforest transformation systems provide increased income and access to education, especially for migrant smallholders. Jungle rubber and rubber monocultures are associated with higher financial land productivity but lower financial labour productivity compared to oil palm, which influences crop choice: smallholders that are labour-scarce would prefer oil palm while land-scarce smallholders would prefer rubber. Collecting long-term data in an interdisciplinary context enables us to provide decision-makers and stakeholders with scientific insights to facilitate the reconciliation between economic interests and ecological sustainability in tropical agricultural landscapes.

Conversion of tropical lowland forest reduces nutrient return through litterfall, and alters nutrient use efficiency and seasonality of net primary production.

Tropical landscapes are not only rapidly transformed by ongoing land-use change, but are additionally confronted by increasing seasonal climate variation. There is an increasing demand for studies analyzing the effects and feedbacks on ecosystem functioning of large-scale conversions of tropical natural forest into intensively managed cash crop agriculture. We analyzed the seasonality of aboveground litterfall, fine root litter production, and aboveground woody biomass production (ANPP(woody)) in natural lowland forests, rubber agroforests under natural tree cover ("jungle rubber"), rubber and oil palm monocultures along a forest-to-agriculture transformation gradient in Sumatra. We hypothesized that the temporal fluctuation of litter production increases with increasing land-use intensity, while the associated nutrient fluxes and nutrient use efficiency (NUE) decrease. Indeed, the seasonal variation of aboveground litter production and ANPP(woody) increased from the natural forest to the plantations, while aboveground litterfall generally decreased. Nutrient return through aboveground litter was mostly highest in the natural forest; however, it was significantly lower only in rubber plantations. NUE of N, P and K was lowest in the oil palm plantations, with natural forest and the rubber systems showing comparably high values. Root litter production was generally lower than leaf litter production in all systems, while the root-to-leaf ratio of litter C flux increased along the land-use intensity gradient. Our results suggest that nutrient and C cycles are more directly affected by climate seasonality in species-poor agricultural systems than in species-rich forests, and therefore might be more susceptible to inter-annual climate fluctuation and climate change.

Quantifying above- and belowground biomass carbon loss with forest conversion in tropical lowlands of Sumatra (Indonesia).

Natural forests in South-East Asia have been extensively converted into other land-use systems in the past decades and still show high deforestation rates. Historically, lowland forests have been converted into rubber forests, but more recently, the dominant conversion is into oil palm plantations. While it is expected that the large-scale conversion has strong effects on the carbon cycle, detailed studies quantifying carbon pools and total net primary production (NPPtotal ) in above- and belowground tree biomass in land-use systems replacing rainforest (incl. oil palm plantations) are rare so far. We measured above- and belowground carbon pools in tree biomass together with NPPtotal in natural old-growth forests, 'jungle rubber' agroforests under natural tree cover, and rubber and oil palm monocultures in Sumatra. In total, 32 stands (eight plot replicates per land-use system) were studied in two different regions. Total tree biomass in the natural forest (mean: 384 Mg ha(-1) ) was more than two times higher than in jungle rubber stands (147 Mg ha(-1) ) and >four times higher than in monoculture rubber and oil palm plantations (78 and 50 Mg ha(-1) ). NPPtotal was higher in the natural forest (24 Mg ha(-1)  yr(-1) ) than in the rubber systems (20 and 15 Mg ha(-1)  yr(-1) ), but was highest in the oil palm system (33 Mg ha(-1)  yr(-1) ) due to very high fruit production (15-20 Mg ha(-1)  yr(-1) ). NPPtotal was dominated in all systems by aboveground production, but belowground productivity was significantly higher in the natural forest and jungle rubber than in plantations. We conclude that conversion of natural lowland forest into different agricultural systems leads to a strong reduction not only in the biomass carbon pool (up to 166 Mg C ha(-1) ) but also in carbon sequestration as carbon residence time (i.e. biomass-C:NPP-C) was 3-10 times higher in the natural forest than in rubber and oil palm plantations.

Patterns in hydraulic architecture from roots to branches in six tropical tree species from cacao agroforestry and their relation to wood density and stem growth.

For decades it has been assumed that the largest vessels are generally found in roots and that vessel size and corresponding sapwood area-specific hydraulic conductivity are acropetally decreasing toward the distal twigs. However, recent studies from the perhumid tropics revealed a hump-shaped vessel size distribution. Worldwide tropical perhumid forests are extensively replaced by agroforestry systems often using introduced species of various biogeographical and climatic origins. Nonetheless, it is unknown so far what kind of hydraulic architectural patterns are developed in those agroforestry tree species and which impact this exerts regarding important tree functional traits, such as stem growth, hydraulic efficiency and wood density (WD). We investigated wood anatomical and hydraulic properties of the root, stem and branch wood in Theobroma cacao and five common shade tree species in agroforestry systems on Sulawesi (Indonesia); three of these were strictly perhumid tree species, and the other three tree species are tolerating seasonal drought. The overall goal of our study was to relate these properties to stem growth and other tree functional traits such as foliar nitrogen content and sapwood to leaf area ratio. Our results confirmed a hump-shaped vessel size distribution in nearly all species. Drought-adapted species showed divergent patterns of hydraulic conductivity, vessel density, and relative vessel lumen area between root, stem and branch wood compared to wet forest species. Confirming findings from natural old-growth forests in the same region, WD showed no relationship to specific conductivity. Overall, aboveground growth performance was better predicted by specific hydraulic conductivity than by foliar traits and WD. Our study results suggest that future research on conceptual trade-offs of tree hydraulic architecture should consider biogeographical patterns underlining the importance of anatomical adaptation mechanisms to environment.