Yield and related traits of three legume crops grown in olive-based agroforestry under an intense drought in the South Mediterranean.

Heat and drought stresses have become more frequent and intense in the Mediterranean, strongly influencing arable crop phenology, growth, and grain yield. Agroforestry systems can effectively buffer the adverse climate conditions and stabilize or even increase crop yield under climate change. However, the positive effects of agroforestry remain uncertain due to the possible intense competition between trees and crops, especially for legume crops that have been less studied than cereals in such context. This study aimed to assess the response of the phenology, growth, grain yield, and yield-related traits of chickpea (Cicer arietinum), faba bean (Vicia faba), and lentil (Lens culinaris) to olive-based agroforestry (AFS) as compared to sole crops system (SCS) in the South of the Mediterranean. We conducted a field experiment during two growing seasons marked by an intense drought, either at the beginning (year 1) or at the end (year 2) of the crop cycle. Crop growth and yield were lower in year 1 than in year 2, reflecting the adverse growing conditions caused by the early drought. They were also lower in AFS than in SCS for both years, indicating that trees had competitive effects on crops. In year 1, the yield loss of grains in AFS was 66 % for lentil, 47 % for chickpea, and 43 % for faba bean compared to SCS, confirming the greater shade sensitivity of lentil. In year 2, the reduction was significantly smaller and was about 46 %, 34 %, 38 % for lentil, chickpea and, faba bean, respectively. The number of pods and grains were the most affected yield components by agroforestry and drought timing across the three legumes crops. Similar responses were found when comparing crops at different distances to trees within the AFS field. Crops generally had lower biomass and yield, explained by fewer pods and grains, on the northern side of trees compared to the southern side of trees or the middle of tree inter-rows, causing significant spatial heterogeneity in crops. However, lentil and chickpea had a positive response to shade during the early drought year while a negative response during the late drought year, suggesting that the benefits of the microclimate created by olive trees express depending on drought timing and crop physiology. Our study supports legume integration into AFS, suggesting that chickpea should be considered during high-stress conditions, while faba bean should be preferred during low-stress conditions.

Drought survival and recovery in grasses: Stress intensity and plant-plant interactions impact plant dehydration tolerance.

Plant dehydration tolerance confers drought survival in grasses, but the mortality thresholds according to soil water content (SWC), vapour pressure deficit (VPD) and plant-plant interactions are little explored. We compared the dehydration dynamics of leaf meristems, which are the key surviving organs, plant mortality, and recovery of Mediterranean and temperate populations of two perennial grass species, Dactylis glomerata and Festuca arundinacea, grown in monocultures and mixtures under a low-VPD (1.5 kPa) versus a high-VPD drought (2.2 kPa). The lethal drought index (LD50 ), that is, SWC associated with 50% plant mortality, ranged from 2.87% (ψs = -1.68 MPa) to 2.19% (ψs = -4.47 MPa) and reached the lowest values under the low-VPD drought. Populations of D. glomerata were more dehydration-tolerant (lower LD50 ), survived and recovered better than F. arundinacea populations. Plant-plant interactions modified dehydration tolerance and improved post-drought recovery in mixtures compared with monocultures. Water content as low as 20.7%-36.1% in leaf meristems allowed 50% of plants to survive. We conclude that meristem dehydration causes plant mortality and that drought acclimation can increase dehydration tolerance. Genetic diversity, acclimation and plant-plant interactions are essential sources of dehydration tolerance variability to consider when predicting drought-induced mortality.

Is a seasonally reduced growth potential a convergent strategy to survive drought and frost in plants?

BACKGROUND: Plants have adapted to survive seasonal life-threatening frost and drought. However, the timing and frequency of such events are impacted by climate change, jeopardizing plant survival. Understanding better the strategies of survival to dehydration stress is therefore timely and can be enhanced by the cross-fertilization of research between disciplines (ecology, physiology), models (woody, herbaceous species) and types of stress (drought, frost). SCOPE: We build upon the 'growth-stress survival' trade-off, which underpins the identification of global plant strategies across environments along a 'fast-slow' economics spectrum. Although phenological adaptations such as dormancy are crucial to survive stress, plant global strategies along the fast-slow economic spectrum rarely integrate growth variations across seasons. We argue that the growth-stress survival trade-off can be a useful framework to identify convergent plant ecophysiological strategies to survive both frost and drought. We review evidence that reduced physiological activity, embolism resistance and dehydration tolerance of meristematic tissues are interdependent strategies that determine thresholds of mortality among plants under severe frost and drought. We show that complete dormancy, i.e. programmed growth cessation, before stress occurrence, minimizes water flows and maximizes dehydration tolerance during seasonal life-threatening stresses. We propose that incomplete dormancy, i.e. the programmed reduction of growth potential during the harshest seasons, could be an overlooked but major adaptation across plants. Quantifying stress survival in a range of non-dormant versus winter- or summer-dormant plants, should reveal to what extent incomplete to complete dormancy could represent a proxy for dehydration tolerance and stress survival. CONCLUSIONS: Our review of the strategies involved in dehydration stress survival suggests that winter and summer dormancy are insufficiently acknowledged as plant ecological strategies. Incorporating a seasonal fast-slow economics spectrum into global plant strategies improves our understanding of plant resilience to seasonal stress and refines our prevision of plant adaptation to extreme climatic events.

Olive agroforestry shapes rhizosphere microbiome networks associated with annual crops and impacts the biomass production under low-rainfed conditions.

Agroforestry (AF) is a promising land-use system to mitigate water deficiency, particularly in semi-arid areas. However, the belowground microbes associated with crops below trees remain seldom addressed. This study aimed at elucidating the effects of olive AF system intercropped with durum wheat (Dw), barely (Ba), chickpea (Cp), or faba bean (Fb) on crops biomass and their soil-rhizosphere microbial networks as compared to conventional full sun cropping (SC) under rainfed conditions. To test the hypothesis, we compared the prokaryotic and the fungal communities inhabiting the rhizosphere of two cereals and legumes grown either in AF or SC. We determined the most suitable annual crop species in AF under low-rainfed conditions. Moreover, to deepen our understanding of the rhizosphere network dynamics of annual crops under AF and SC systems, we characterized the microbial hubs that are most likely responsible for modifying the microbial community structure and the variability of crop biomass of each species. Herein, we found that cereals produced significantly more above-ground biomass than legumes following in descending order: Ba > Dw > Cp > Fb, suggesting that crop species play a significant role in improving soil water use and that cereals are well-suited to rainfed conditions within both types of agrosystems. The type of agrosystem shapes crop microbiomes with the only marginal influence of host selection. However, more relevant was to unveil those crops recruits specific bacterial and fungal taxa from the olive-belowground communities. Of the selected soil physicochemical properties, organic matter was the principal driver in shaping the soil microbial structure in the AF system. The co-occurrence network analyses indicated that the AF system generates higher ecological stability than the SC system under stressful climate conditions. Furthermore, legumes' rhizosphere microbiome possessed a higher resilient capacity than cereals. We also identified different fungal keystones involved in litter decomposition and drought tolerance within AF systems facing the water-scarce condition and promoting crop production within the SC system. Overall, we showed that AF reduces cereal and legume rhizosphere microbial diversity, enhances network complexity, and leads to more stable beneficial microbial communities, especially in severe drought, thus providing more accurate predictions to preserve soil diversity under unfavorable environmental conditions.

Can trees buffer the impact of climate change on pasture production and digestibility of Mediterranean dehesas?

Sustainability and functioning of silvopastoral ecosystems are being threatened by the forecasted warmer and drier environments in the Mediterranean region. Scattered trees of these ecosystems could potentially mitigate the impact of climate change on herbaceous plant community but this issue has not yet tested experimentally. We carried out a field manipulative experiment of increased temperature (+2-3 °C) using Open Top Chambers and rainfall reduction (30%) through rain-exclusion shelters to evaluate how net primary productivity and digestibility respond to climate change over three consecutive years, and to test whether scattered trees could buffer the effects of higher aridity in Mediterranean dehesas. First, we observed that herbaceous communities located beneath tree canopy were less productive (351 g/m2) than in open grassland (493 g/m2) but had a higher digestibility (44% and 41%, respectively), likely promoted by tree shade and the higher soil fertility of this habitat. Second, both habitats responded similarly to climate change in terms of net primary productivity, with a 33% increase under warming and a 13% decrease under reduced rainfall. In contrast, biomass digestibility decreased under increased temperatures (-7.5%), since warming enhanced the fiber and lignin content and decreased the crude protein content of aerial biomass. This warming-induced effect on biomass digestibility only occurred in open grasslands, suggesting a buffering role of trees in mitigating the impact of climate change. Third, warming did not only affect these ecosystem processes in a direct way but also indirectly via changes in plant functional composition. Our findings suggest that climate change will alter both the quantity and quality of pasture production, with expected warmer conditions increasing net primary productivity but at the expense of reducing digestibility. This negative effect of warming on digestibility might be mitigated by scattered trees, highlighting the importance of implementing strategies and suitable management to control tree density in these ecosystems.

Research on agroforestry systems and biodiversity conservation: what can we conclude so far and what should we improve?

Through a meta-analysis, Mupepele et al. (BMC Ecol Evol 21:1-193, 2021) assessed the effects of European agroforestry systems on biodiversity, estimated by species richness or species diversity. They showed that the effects of silvoarable and silvopastoral systems depend on the systems they are compared to and the taxa studied. Further, they found that only silvoarable systems increased species richness or diversity, compared to cropland. The authors conclude that agroforestry systems have weak effects on biodiversity and that landscape context or land-use history are probably more important than the practice of agroforestry in itself. However, we draw attention to important shortcomings in this meta-analysis, which downplay the potential of agroforestry for biodiversity conservation in agricultural landscapes. We hope that the meta-analysis by Mupepele et al. (BMC Ecol Evol 21:1-193, 2021), and our comments, will contribute to improving the quality of research on agroforestry systems and biodiversity conservation.