Plant root mechanisms and their effects on carbon and nutrient accumulation in desert ecosystems under changes in land use and climate.

Deserts represent key carbon reservoirs, yet as these systems are threatened this has implications for biodiversity and climate change. This review focuses on how these changes affect desert ecosystems, particularly plant root systems and their impact on carbon and mineral nutrient stocks. Desert plants have diverse root architectures shaped by water acquisition strategies, affecting plant biomass and overall carbon and nutrient stocks. Climate change can disrupt desert plant communities, with droughts impacting both shallow and deep-rooted plants as groundwater levels fluctuate. Vegetation management practices, like grazing, significantly influence plant communities, soil composition, root microorganisms, biomass, and nutrient stocks. Shallow-rooted plants are particularly susceptible to climate change and human interference. To safeguard desert ecosystems, understanding root architecture and deep soil layers is crucial. Implementing strategic management practices such as reducing grazing pressure, maintaining moderate harvesting levels, and adopting moderate fertilization can help preserve plant-soil systems. Employing socio-ecological approaches for community restoration enhances carbon and nutrient retention, limits desert expansion, and reduces CO2 emissions. This review underscores the importance of investigating belowground plant processes and their role in shaping desert landscapes, emphasizing the urgent need for a comprehensive understanding of desert ecosystems.

Impact of aridity rise and arid lands expansion on carbon-storing capacity, biodiversity loss, and ecosystem services.

Drylands, comprising semi-arid, arid, and hyperarid regions, cover approximately 41% of the Earth's land surface and have expanded considerably in recent decades. Even under more optimistic scenarios, such as limiting global temperature rise to 1.5°C by 2100, semi-arid lands may increase by up to 38%. This study provides an overview of the state-of-the-art regarding changing aridity in arid regions, with a specific focus on its effects on the accumulation and availability of carbon (C), nitrogen (N), and phosphorus (P) in plant-soil systems. Additionally, we summarized the impacts of rising aridity on biodiversity, service provisioning, and feedback effects on climate change across scales. The expansion of arid ecosystems is linked to a decline in C and nutrient stocks, plant community biomass and diversity, thereby diminishing the capacity for recovery and maintaining adequate water-use efficiency by plants and microbes. Prolonged drought led to a -3.3% reduction in soil organic carbon (SOC) content (based on 148 drought-manipulation studies), a -8.7% decrease in plant litter input, a -13.0% decline in absolute litter decomposition, and a -5.7% decrease in litter decomposition rate. Moreover, a substantial positive feedback loop with global warming exists, primarily due to increased albedo. The loss of critical ecosystem services, including food production capacity and water resources, poses a severe challenge to the inhabitants of these regions. Increased aridity reduces SOC, nutrient, and water content. Aridity expansion and intensification exacerbate socio-economic disparities between economically rich and least developed countries, with significant opportunities for improvement through substantial investments in infrastructure and technology. By 2100, half the world's landmass may become dryland, characterized by severe conditions marked by limited C, N, and P resources, water scarcity, and substantial loss of native species biodiversity. These conditions pose formidable challenges for maintaining essential services, impacting human well-being and raising complex global and regional socio-political challenges.

Combining different species in restoration is not always the right decision: Monocultures can provide higher ecological functions than intercropping in a desert ecosystem.

Vegetation restoration in deserts is challenging due to these ecosystems' inherent fragility and harsh environmental conditions. One approach for active restoration involves planting native species, which can accelerate the recovery of ecosystem functions. To ensure the effectiveness of this process, carefully selecting species for planting is crucial. Generally, it is expected that a more diverse mix of species in the plantation will lead to the recovery of a greater number of ecosystem functions, especially when the selected species have complementary niche traits that facilitate maximum cooperation and minimize competition among them. In this study, we evaluated the planting of two native species from the hyper-desert of Taklamakan, China, which exhibit marked morpho-physiological differences: a phreatophytic legume (Alhagi sparsifolia) and a halophytic non-legume (Karelinia caspia). These species were grown in both monoculture and intercrop communities. Monoculture of the legume resulted in the highest biomass accumulation. Intercropping improved several ecosystem functions in the 50 cm-upper soil, particularly those related to phosphorus (P), carbon (C), and sulfur (S) concentrations, as well as soil enzyme activities. However, it also increased soil sodium (Na+) concentration and pH. Halophyte monocultures enhanced ecological functions associated with nitrogen concentrations in the upper soil and with P, S, C, and cation concentrations (K+, Ca2+, Mg2+, Cu2+, Fe2+, Zn2+, Co2+, Ni2+), along with enzyme activities in the deep soil. It also maximized Na+ accumulation in plant biomass. In summary, we recommend legume monoculture when the primary goal is to optimize biomass accumulation. Conversely, halophyte monoculture is advisable when the objective is to extract sodium from the soil or enhance ecosystem functions in the deep soil. Intercropping the two species is recommended to maximize the ecosystem functions of the upper soil, provided there is no salinization risk. When planning restoration efforts in desert regions, it is essential to understand the impact of each species on ecosystem function and how complementary species behave when intercropped. However, these interactions are likely species- and system-specific, highlighting the need for more work to optimize solutions for different arid ecosystems.

Strigolactones can be a potential tool to fight environmental stresses in arid lands.

BACKGROUND: Environmental stresses pose a significant threat to plant growth and ecosystem productivity, particularly in arid lands that are more susceptible to climate change. Strigolactones (SLs), carotenoid-derived plant hormones, have emerged as a potential tool for mitigating environmental stresses. METHODS: This review aimed to gather information on SLs' role in enhancing plant tolerance to ecological stresses and their possible use in improving the resistance mechanisms of arid land plant species to intense aridity in the face of climate change. RESULTS: Roots exude SLs under different environmental stresses, including macronutrient deficiency, especially phosphorus (P), which facilitates a symbiotic association with arbuscular mycorrhiza fungi (AMF). SLs, in association with AMF, improve root system architecture, nutrient acquisition, water uptake, stomatal conductance, antioxidant mechanisms, morphological traits, and overall stress tolerance in plants. Transcriptomic analysis revealed that SL-mediated acclimatization to abiotic stresses involves multiple hormonal pathways, including abscisic acid (ABA), cytokinins (CK), gibberellic acid (GA), and auxin. However, most of the experiments have been conducted on crops, and little attention has been paid to the dominant vegetation in arid lands that plays a crucial role in reducing soil erosion, desertification, and land degradation. All the environmental gradients (nutrient starvation, drought, salinity, and temperature) that trigger SL biosynthesis/exudation prevail in arid regions. The above-mentioned functions of SLs can potentially be used to improve vegetation restoration and sustainable agriculture. CONCLUSIONS: Present review concluded that knowledge on SL-mediated tolerance in plants is developed, but still in-depth research is needed on downstream signaling components in plants, SL molecular mechanisms and physiological interactions, efficient methods of synthetic SLs production, and their effective application in field conditions. This review also invites researchers to explore the possible application of SLs in improving the survival rate of indigenous vegetation in arid lands, which can potentially help combat land degradation problems.

Alhagi sparsifolia acclimatizes to saline stress by regulating its osmotic, antioxidant, and nitrogen assimilation potential.

BACKGROUND: Alhagi sparsifolia (Camelthorn) is a leguminous shrub species that dominates the Taklimakan desert's salty, hyperarid, and infertile landscapes in northwest China. Although this plant can colonize and spread in very saline soils, how it adapts to saline stress in the seedling stage remains unclear so a pot-based experiment was carried out to evaluate the effects of four different saline stress levels (0, 50, 150, and 300 mM) on the morphological and physio-biochemical responses in A. sparsifolia seedlings. RESULTS: Our results revealed that N-fixing A. sparsifolia has a variety of physio-biochemical anti-saline stress acclimations, including osmotic adjustments, enzymatic mechanisms, and the allocation of metabolic resources. Shoot-root growth and chlorophyll pigments significantly decreased under intermediate and high saline stress. Additionally, increasing levels of saline stress significantly increased Na+ but decreased K+ concentrations in roots and leaves, resulting in a decreased K+/Na+ ratio and leaves accumulated more Na + and K + ions than roots, highlighting their ability to increase cellular osmolarity, favouring water fluxes from soil to leaves. Salt-induced higher lipid peroxidation significantly triggered antioxidant enzymes, both for mass-scavenging (catalase) and cytosolic fine-regulation (superoxide dismutase and peroxidase) of H2O2. Nitrate reductase and glutamine synthetase/glutamate synthase also increased at low and intermediate saline stress levels but decreased under higher stress levels. Soluble proteins and proline rose at all salt levels, whereas soluble sugars increased only at low and medium stress. The results show that when under low-to-intermediate saline stress, seedlings invest more energy in osmotic adjustments but shift their investment towards antioxidant defense mechanisms under high levels of saline stress. CONCLUSIONS: Overall, our results suggest that A. sparsifolia seedlings tolerate low, intermediate, and high salt stress by promoting high antioxidant mechanisms, osmolytes accumulations, and the maintenance of mineral N assimilation. However, a gradual decline in growth with increasing salt levels could be attributed to the diversion of energy from growth to maintain salinity homeostasis and anti-stress oxidative mechanisms.

Impact of phosphorus application on drought resistant responses of Eucalyptus grandis seedlings.

Eucalyptus grandis is the most widely planted tree species worldwide and can face severe drought during the initial months after planting because the root system is developing. A complete randomized design was used to study the effects of two water regimes (well-watered and water-stressed) and phosphorus (P) applications (with and without P) on the morphological and physio-biochemical responses of E. grandis. Drought had negative effects on the growth and metabolism of E. grandis, as indicated by changes in morphological traits, decreased net photosynthetic rates (Pn ), pigment concentrations, leaf relative water contents (LRWCs), nitrogenous compounds, over-production of reactive oxygen species (ROS) and higher lipid peroxidation. However, E. grandis showed effective drought tolerance strategies, such as reduced leaf area and transpiration rate (E), higher accumulation of soluble sugars and proline and a strong antioxidative enzyme system. P fertilization had positive effects on well-watered seedlings due to improved growth and photosynthesis, which indicated the high P requirements during the initial E. grandis growth stage. In drought-stressed seedlings, P application had no effects on the morphological traits, but it significantly improved the LRWC, Pn , quantum efficiency of photosystem II (Fv /Fm ), chlorophyll pigments, nitrogenous compounds and reduced lipid peroxidation. P fertilization improved E. grandis seedling growth under well-watered conditions but also ameliorated some leaf physiological traits under drought conditions. The effects of P fertilization are mainly due to the enhancement of plant N nutrition. Therefore, P can be used as a fertilizer to improve growth and production in the face of future climate change.

Soil microbial functional profiles of P-cycling reveal drought-induced constraints on P-transformation in a hyper-arid desert ecosystem.

Soil water conditions are known to influence soil nutrient availability, but the specific impact of different conditions on soil phosphorus (P) availability through the modulation of P-cycling functional microbial communities in hyper-arid desert ecosystems remains largely unexplored. To address this knowledge gap, we conducted a 3-year pot experiment using a typical desert plant species (Alhagi sparsifolia Shap.) subjected to two water supply levels (25 %-35 % and 65 %-75 % of maximum field capacity, MFC) and four P-supply levels (0, 1, 3, and 5 g P m-2 y-1). Our investigation focused on the soil Hedley-P pool and the four major microbial groups involved in the critical phases of soil microbial P-cycling. The results revealed that the drought (25 %-35 % MFC) and no P-supply treatments reduced soil resin-P and NaHCO3-Pi concentrations by 87.03 % and 93.22 %, respectively, compared to the well-watered (65 %-75 % MFC) and high P-supply (5 g P m-2 y-1) treatments. However, the P-supply treatment resulted in a 12 %-22 % decrease in the soil NH4+-N concentration preferred by microbes compared to the no P-supply treatment. Moreover, the abundance of genes engaged in microbial P-cycling (e.g. gcd and phoD) increased under the drought and no P-supply treatments (p < 0.05), suggesting that increased NH4+-N accumulation under these conditions may stimulate P-solubilizing microbes, thereby promoting the microbial community's investment in resources to enhance the P-cycling potential. Furthermore, the communities of Steroidobacter cummioxidans, Mesorhizobium alhagi, Devosia geojensis, and Ensifer sojae, associated with the major P-cycling genes, were enriched in drought and no or low-P soils. Overall, the drought and no or low-P treatments stimulated microbial communities and gene abundances involved in P-cycling. However, this increase was insufficient to maintain soil P-bioavailability. These findings shed light on the responses and feedback of microbial-mediated P-cycling behaviors in desert ecosystems under three-year drought and soil P-deficiency.

Intercropping of Leguminous and Non-Leguminous Desert Plant Species Does Not Facilitate Phosphorus Mineralization and Plant Nutrition.

More efficient use of soil resources, such as nitrogen (N) and phosphorus (P), can improve plant community resistance and resilience against drought in arid and semi-arid lands. Intercropping of legume and non-legumes can be an effective practice for enhancing P mineralization uptake, and plant nutrient status. However, it remains unclear how intercropping systems using desert plant species impact soil-plant P fractions and how they affect N and water uptake capacity. Alhagi sparsifolia (a legume) and Karelinia caspia (a non-legume) are dominant plant species in the Taklamakan Desert in Xinjiang Province, China. However, there is a lack of knowledge of whether these species, when intercropped, can trigger synergistic processes and mechanisms that drive more efficient use of soil resources. Thus, in a field experiment over two years, we investigated the impact of monoculture and intercropping of these plant species on soil-plant P fractions and soil-plant nutrients. Both plant species' foliar nutrient (N, P, and K) concentrations were higher under monoculture than intercropping (except K in K. caspia). Nucleic acid P was higher in the monoculture plots of A. sparsifolia, consistent with higher soil labile P, while metabolic P was higher in monoculture K. caspia, associated with higher soil moderately labile Pi. However, both species had a higher residual P percentage in the intercropping system. Soils from monoculture and intercropped plots contained similar microbial biomass carbon (MBC), but lower microbial biomass N:microbial biomass phosphorus (MBN:MBP) ratio associated with reduced N-acetylglucosaminidase (NAG) activity in the intercropped soils. This, together with the high MBC:MBN ratio in intercropping and the lack of apparent general effects of intercropping on MBC:MBP, strongly suggest that intercropping improved microbe N- but not P-use efficiency. Interestingly, while EC and SWC were higher in the soil of the K. caspia monoculture plots, EC was significantly lower in the intercropped plots. Plants obtained better foliar nutrition and soil P mineralization in monocultures than in intercropping systems. The possible positive implications of intercropping for reducing soil salinization and improving soil water uptake and microbial N-use efficiency could have advantages in the long term and its utilization should be explored further in future studies.

Phosphorous Supplementation Alleviates Drought-Induced Physio-Biochemical Damages in Calligonum mongolicum.

Calligonum mongolicum is a phreatophyte playing an important role in sand dune fixation, but little is known about its responses to drought and P fertilization. In the present study, we performed a pot experiment to investigate the effects of P fertilization under drought or well-watered conditions on multiple morpho-physio-biochemical attributes of C. mongolicum seedlings. Drought stress leads to a higher production of hydrogen peroxide (H2O2) and malondialdehyde (MDA), leading to impaired growth and metabolism. However, C. mongolicum exhibited effective drought tolerance strategies, including a higher accumulation of soluble sugars, starch, soluble protein, proline, and significantly higheractivities of peroxidase (POD) and catalase (CAT) enzymes. P fertilization increased the productivity of drought-stressed seedlings by increasing their growth, assimilative shoots relative water content, photosynthetic pigments, osmolytes accumulation, mineral nutrition, N assimilation, and reduced lipid peroxidation. Our findings suggest the presence of soil high P depletion and C. mongolicum high P requirements during the initial growth stage. Thus, P can be utilized as a fertilizer to enhance the growth and productivity of Calligonum vegetation and to reduce the fragility of the hyper-arid desert of Taklamakan in the context of future climate change.

Alhagi sparsifolia: An ideal phreatophyte for combating desertification and land degradation.

The upcoming launch of the post-2020 biodiversity framework entails elucidating the future priorities and strategies for conserving biodiversity on a regional and global scale. Desertification has caused significant environmental and socio-economic problems in many arid and semiarid areas of the world. Phreatophytes increase nutrient levels and water reserves in the ground below them and are essential resources for other plants and animals. Factors that can enhance or reduce the island effect of phreatophytes are worthy of research since they can contribute to the survival and growth of this group of plants. Furthermore, these plants substantially impact the whole environment and boast necessary evolutionary adaptations for sustaining islands of life in the arid landscape to prevent complete desertification. Alhagi sparsifolia is a semi-shrub legume and a typical desert phreatophyte. The main objective of this review is to disseminate research-based knowledge and improve awareness of this important plant species and provide a framework that could effectively improve regional socio-ecological environments, particularly in controlling desertification, thereby enhancing human welfare. First, we synthesized knowledge of the physiological acclimation of this species, above all in terms of water and nutrient use, the impact of salinity, and its response to different light environments and temperatures. Then, we analyzed the socio-ecological services that A. sparsifolia offers, highlighted the anthropogenic threats to its habitats, and strategies for its sustainable utilization and management. We conclude that more knowledge of how A. sparsifolia interacts with groundwater will contribute significantly to elucidating the possible effects of climate change and human activities on the sustainable use of groundwater. Protected areas must be set up, and key knowledge shared to combat desertification and improve people's welfare as a means of guaranteeing sustainable development. An integrative socio-ecological approach must be adapted to improve understanding of this valuable phreatophyte's importance and sustainable use and fulfill Sustainable Development Goals (SDGs).

Stoichiometry of C:N:P in the Roots of Alhagi sparsifolia Is More Sensitive to Soil Nutrients Than Aboveground Organs.

The stoichiometry of carbon, nitrogen, and phosphorus (C:N:P) among leaves, stems, and roots reflects trade-offs in plants for acquiring resources and their growth strategy. The widely distributed plant Alhagi sparsifolia is an ideal species to study the ecological stoichiometry in different organs in response to the availability of nutrients and water in the desert ecosystem. However, which response of organs is most sensitive to environmental conditions is still unclear. To answer this question, we collected samples of plants and soils including not only aboveground leaves and stems, but also underground roots and soils from a wide range of arid areas during the growing season. The C, N, P, C:N, C:P, and N:P ratios in leaves, thorns, stems, and roots were derived to explore their relationship as well as their response mechanisms to nutrients and water spanning 1 m deep in the soil. The results showed that the order of N concentration was leaves > thorns > stems > roots, that the concentration of P in the leaves, thorns, and stems was similar, and that their values were higher than those in the roots. First, the C:N ratios in the leaves and stems were significantly positively correlated with the ratio in roots. The C:N ratios in each organ showed a significant relationship with the soil alkali hydrolyzable nitrogen (SAN) above a depth of 60 cm. In addition to SAN, soil available phosphorus (SAP) and soil organic carbon (SOC) affect the C:N ratio in the roots. Second, the C:P and N:P ratios in aboveground organs showed no correlations with the ratios in roots. The C:P and N:P ratios in the leaves and thorns have no relationship with soil nutrients, while the C:P ratio in roots was influenced by SAN and SOC in all soil layers. Finally, the N:P ratios in roots were also affected by nutrients in different soil depths at 0-20 and 60-80 cm. These results illustrate that the roots were more sensitive to soil nutrients than the aboveground parts. Our study of ecological stoichiometry also suggests a novel systematic approach for analyzing the sensitivity of responses of an organ to environmental conditions.

Intercropping Systems Modify Desert Plant-Associated Microbial Communities and Weaken Host Effects in a Hyper-Arid Desert.

Intercropping is an important practice in promoting plant diversity and productivity. Compared to the accumulated understanding of the legume/non-legume crop intercrops, very little is known about the effect of this practice when applied to native species on soil microbial communities in the desert ecosystem. Therefore, in the present study, bulk soil and rhizosphere microbial communities in the 2-year Alhagi sparsifolia (legume)/Karelinia caspica (non-legume) monoculture vs. intercropping systems were characterized under field conditions. Our result revealed that plant species identities caused a significant effect on microbial community composition in monocultures but not in intercropping systems. Monoculture weakened the rhizosphere effect on fungal richness. The composition of bacterial and fungal communities (ß-diversity) was significantly modified by intercropping, while bacterial richness (Chao1) was comparable between the two planting patterns. Network analysis revealed that Actinobacteria, α- and γ-proteobacteria dominated bulk soil and rhizosphere microbial co-occurrence networks in each planting pattern. Intercropping systems induced a more complex rhizosphere microbial community and a more modular and stable bulk soil microbial network. Keystone taxa prevailed in intercropping systems and were Actinobacteria-dominated. Overall, planting patterns and soil compartments, not plant identities, differentiated root-associated microbiomes. Intercropping can modify the co-occurrence patterns of bulk soil and rhizosphere microorganisms in desert ecosystems. These findings provided a potential strategy for us to manipulate desert soil microbial communities and optimize desert species allocation in vegetation sustainability.

Coupling Relationship of Leaf Economic and Hydraulic Traits of Alhagisparsifolia Shap. in a Hyper-Arid Desert Ecosystem.

In this study, Alhagisparsifolia Shap. was used to test the hypothesis that leaf economic and hydraulic traits are coupled in plants in a hyper-arid region. Five economic traits and six hydraulic traits were examined to explore the relationship. Results showed that the stomatal density (SD) on both surfaces was coupled with maximum stomatal conductance to water vapor (gwmax) and leaf tissue density (TD). SD on adaxial surface (SDaba) was significantly positively related to vein density (VD) but negatively related to leaf thickness (LT) and stomatal length on adaxial surface (SLada). Nitrogen concentration based on mass (Nmass) was significantly negatively correlated with leaf mass per area (LMA), LT, and VD, whereas nitrogen concentration based on area (Narea) was significantly positively related to LMA and TD. Mean annual precipitation (MAP) contributed the most to the changes in LT and stomatal length (SL). Soil salt contributed the most to TD, SD, and gwmax. Soli nutrients influenced the most of LMA and VD. Mean annual temperature contributed the most to Nmass and Narea. In conclusion, the economics of leaves coupled with their hydraulic traits provides an economical and efficient strategy to adapt to the harsh environment in hyper-arid regions.

Nitrogen application mitigates drought-induced metabolic changes in Alhagi sparsifolia seedlings by regulating nutrient and biomass allocation patterns.

Groundwater and its associated nutrients sustain the establishment and persistence of phreatophytes. Rapid root elongation immediately after germination is vital for desert species to access deep water sources to avoid water-deficit stress. However, the growth strategy and responses to nutrients and water of young phreatophyte seedlings before their roots reach the water table are poorly understood, especially in the scenarios of nitrogen (N) deposition and drought. We investigated how simulated N deposition and drought affect the plasticity of Alhagi sparsifolia seedlings by multiple eco-physiological mechanisms. Seedlings were planted under drought-stressed or well-watered conditions and subjected to various levels of N addition (0, 3.0, 6.0, or 9.0 gN·m-2 yr-1). The amounts of N and water independently or interactively affected the photosynthetic traits, drought tolerance characteristics, morphological traits, biomass allocation strategy, and nutrient distribution patterns among the plant organs. Moreover, changes mediated by N addition at the leaf level reflected the drought acclimation of the seedlings, which may be related to biomass and nutrient partitioning between organs. The roots were found to be more sensitive to variation of the N:phosphorus (P) ratio, and greater proportions of biomass, N, and P were allocated to resource-acquiring organs (i.e., leaves and fine roots) than to other tissues. A. sparsifolia adopts numerous strategies to tolerate drought, and additional N input was crucial to enhance the growth of drought-stressed A. sparsifolia, which was mainly attributable to its positive impact on the N and P uptake capacity mediated by increased biomass allocation to the roots.

Role of Glycine max in improving drought tolerance in Zanthoxylum bungeanum.

Intercropping may improve community stability and yield under climate change. Here, we set up a field experiment to evaluate the advantages of cultivating Z anthoxylum bungeanum with Capsicum annum, and Z. bungeanum with Glycine max as intercrops, compared with cultivating Z. bungeanum in monoculture. Effects of extreme drought stress conditions on morphological, physiological, and biochemical traits of the three crop species cultivated in the three contrasting planting systems were compared. Results showed that extreme drought conditions induced negative impacts on Z. bungeanum grown in monoculture, due to reduced growth and metabolic impairment. However, limited stomatal conductance, reduced transpiration rate (Tr ), and increased water use efficiency, carotenoid content, catalase activity, and accumulation of soluble sugars in Z. bungeanum indicated its adaptive strategies for tolerance of extreme drought stress conditions. Compared with cultivation in monoculture, intercropping with C. annum had positive effects on Z. bungeanum under extreme drought stress conditions, as a result of improved crown diameter, leaf relative water content (LRWC), net photosynthetic rate, and proline content, while intercropping with G. max under extreme drought stress conditions increased net CO2 assimilation rates, LRWC, Tr , and superoxide dismutase (SOD) activity. In conclusion, Z. bungeanum has an effective defense mechanism for extreme drought stress tolerance. Intercropping with G. max enhanced this tolerance potential primarily through its physio-biochemical adjustments, rather than as a result of nitrogen fixation by G. max.

Growth promotion and protection from drought in Eucalyptus grandis seedlings inoculated with beneficial bacteria embedded in a superabsorbent polymer.

Eucalyptus grandis is a globally important tree crop. Greenhouse-grown tree seedlings often face water deficit after outplanting to the field, which can affect their survival and establishment severely. This can be alleviated by the application of superabsorbent hydrophilic polymers (SAPs). Growth promoting bacteria can also improve crop abiotic stress tolerance; however, their use in trees is limited, partly due to difficulties in the application and viability loss. In this work, we evaluated the improvement of drought tolerance of E. grandis seedlings by inoculating with two Pseudomonas strains (named M25 and N33), carried by an acrylic-hydrocellulosic SAP. We observed significant bacterial survival in the seedling rhizosphere 50 days after inoculation. Under gradual water deficit conditions, we observed a considerable increase in the water content and wall elasticity of M25-inoculated plants and a trend towards growth promotion with both bacteria. Under rapid water deficit conditions, which caused partial defoliation, both strains significantly enhanced the formation of new leaves, while inoculation with M25 reduced the transpiration rate. Co-inoculation with M25 and N33 substantially increased growth and photosynthetic capacity. We conclude that the selected bacteria can benefit E. grandis early growth and can be easily inoculated at transplant by using an acrylic-hydrocellulosic SAP.

Poplar leaf rust reduces dry mass accumulation and internal nitrogen recycling more markedly under low soil nitrogen availability, and decreases growth in the following spring.

Rust is one of the most important biotic stress factors that affect poplars. The aims of this work were: (i) to analyze the changes in growth and nitrogen (N) accumulation in Populus deltoides W. Bartram ex Marshall plants infected with rust (Melampsora medusae Thümen.) and to determine how internal N stores are affected by the disease, in plants growing under two N availabilities in the soil; and (ii) to evaluate the impact of rust in the early sprout in the following growing season and the cumulative effect of the disease after repeated infections. Two clones with different susceptibility to rust were analyzed. At leaf level, rust reduced gas exchange capacity, water conductance in liquid phase and photosynthetic rate in both clones. At plant level, rust reduced plant growth, accelerated leaf senescence and abscission occurred with a higher concentration of leaf N. Even though N concentration in stems and roots were not significantly reduced by rust, total N accumulation in perennial tissues was reduced in infected plants. The vigor of the early sprout of plants infected by rust in the previous season was lower than that of non-infected plants. Therefore, rust affects plant growth by reducing the photosynthetic capacity and leaf area duration, and by decreasing internal nutrient recycling. As nutrient reserves in perennial tissues are lower, rust infection reduces not only the growth of the current season, but also has a cumulative effect on the following years. The reduction of growth was similar in both clones. High availability of N in the soil had no effect on leaf physiology but increased plant growth, delayed leaf senescence and abscission, and increased total N accumulation. If fertilization increases plant growth (stem and root dry mass) it can mitigate the negative effect of the pathogen in the reduction of nutrient storages and future growth.

Plant-pathogen interactions: leaf physiology alterations in poplars infected with rust (Melampsora medusae).

Rust produced by Melampsora sp. is considered one of the most relevant diseases in poplar plantations. Growth reduction in poplar plantations takes place because rust, like other pathogens, alters leaf physiology. There is not a complete evaluation of several of the physiological traits that can be affected by rust at leaf level. Therefore, the aim of this work was to evaluate, in an integrative way and in the same pathosystem, which physiological processes are affected when Populus deltoides Bartr. ex Marsh. leaves are infected by rust (Melampsora medusae Thümen). Leaves of two clones with different susceptibility to rust were analyzed. Field and pot experiments were performed, and several physiological traits were measured in healthy and infected leaves. We conclude that rust affects leaf mesophyll integrity, and so water movement in the leaf in liquid phase is affected. As a consequence, gas exchange is reduced, affecting both carbon fixation and transpiration. However, there is an increase in respiration rate, probably due to plant and fungal respiration. The increase in respiration rate is important in the reduction of net photosynthetic rate, but also some damage in the photosynthetic apparatus limits leaf capacity to fix carbon. The decrease in chlorophyll content would start later and seems not to explain the reduction in net photosynthetic rate. Both clones, although they have different susceptibility to rust, are affected in the same physiological mechanisms.

Phosphorous fertilization alleviates drought effects on Alnus cremastogyne by regulating its antioxidant and osmotic potential.

Alnus cremastogyne, a broad-leaved tree endemic to south-western China, has both commercial and restoration importance. However, little is known of its morphological, physiological and biochemical responses to drought and phosphorous (P) application. A randomized experimental design was used to investigate how drought affected A. cremastogyne seedlings, and the role that P applications play in these responses. Drought had significant negative effects on A. cremastogyne growth and metabolism, as revealed by reduced biomass (leaf, shoot and root), leaf area, stem diameter, plant height, photosynthetic rate, leaf relative water content, and photosynthetic pigments, and a weakened antioxidative defence mechanism and high lipid peroxidation level. However, the reduced leaf area and enhanced osmolyte (proline and soluble sugars) accumulation suggests drought avoidance and tolerance strategies in this tree. Applying P significantly improved the leaf relative water content and photosynthetic rate of drought-stressed seedlings, which may reflect increased anti-oxidative enzyme (superoxide dismutase, catalase and peroxidase) activities, osmolyte accumulation, soluble proteins, and decreased lipid peroxidation levels. However, P had only a slight or negligible effect on the well-watered plants. A. cremastogyne is sensitive to drought stress, but P facilitates and improves its metabolism primarily via biochemical and physiological rather than morphological adjustments, regardless of water availability.

Phosphorous Application Improves Drought Tolerance of Phoebe zhennan.

Phoebe zhennan (Gold Phoebe) is a threatened tree species in China and a valuable and important source of wood and bioactive compounds used in medicine. Apart from anthropogenic disturbances, several biotic constraints currently restrict its growth and development. However, little attention has been given to building adaptive strategies for its conservation by examining its morphological and physio-biochemical responses to drought stress, and the role of fertilizers on these responses. A randomized experimental design was used to investigate the effects of two levels of irrigation (well-watered and drought-stressed) and phosphorous (P) fertilization treatment (with and without P) to assess the morphological and physio-biochemical responses of P. zhennan seedlings to drought stress. In addition, we evaluated whether P application could mitigate the negative impacts of drought on plant growth and metabolism. Drought stress had a significant negative effect on the growth and metabolic processes of P. zhennan. Despite this, reduced leaf area, limited stomatal conductance, reduced transpiration rate, increased water use efficiency, enhanced antioxidant enzymes activities, and osmolytes accumulation suggested that the species has good adaptive strategies for tolerating drought stress. Application of P had a significant positive effect on root biomass, signifying its improved water extracting capacity from the soil. Moreover, P fertilization significantly increased leaf relative water content, net photosynthetic rate, and maximal quantum efficiency of PSII under drought stress conditions. This may be attributable to several factors, such as enhanced root biomass, decreased malondialdehyde content, and the up-regulation of chloroplast pigments, osmolytes, and nitrogenous compounds. However, P application had only a slight or negligible effect on the growth and metabolism of well-watered plants. In conclusion, P. zhennan has a strong capability for drought resistance, while P application facilitates and improves drought tolerance mostly through physio-biochemical adjustments, regardless of water availability. It is imperative to explore the underlying metabolic mechanisms and effects of different levels of P fertilization on P. zhennan under drought conditions in order to design appropriate conservation and management strategies for this species, which is at risk of extinction.