Impact of shrub canopies on understorey vegetation in western Eurasian tundra

Question: How does the composition and species richness of understorey vegetation associate with changing abundance of deciduous shrub canopies? What are the species‐specific associations between shrubs and understorey plants? Location: Tundra habitats along an over 1000‐km long range, spanning from NW Fennoscandia to the Yamal Peninsula in northwest Russia. Methods: The data from 758 vegetation sample plots from 12 sites comprised cover estimates of all plant species, including bryophytes and lichens, and canopy height of deciduous shrubs. The relationships between shrub volume and cover of plant groups and species richness of vegetation were investigated. In addition, species‐specific associations between understorey species and shrub volume were analysed. Results: Shrub abundance was shown to be associated with the composition of understorey vegetation, and the association patterns were consistent across the study sites. Increased forb cover was positively associated with shrub volume, whereas bryophyte, lichen, dwarf shrub and graminoid cover decreased in association with increasing volume of deciduous shrubs. The total species richness of vegetation declined with increasing shrub volume. Conclusions: The results suggest that an increase of shrubs – due to climatic warming or a decrease in grazing pressure – is likely to have strong effects on plant–plant interactions and lead to a decrease in the diversity of understorey vegetation.     

Plant functional group responses to fire frequency and tree canopy cover gradients in oak savannas and woodlands

Questions: How do fire frequency, tree canopy cover, and their interactions influence cover of grasses, forbs and understorey woody plants in oak savannas and woodlands? Location: Minnesota, USA. Methods: We measured plant functional group cover and tree canopy cover on permanent plots within a long‐term prescribed fire frequency experiment and used hierarchical linear modeling to assess plant functional group responses to fire frequency and tree canopy cover. Results: Understorey woody plant cover was highest in unburned woodlands and was negatively correlated with fire frequency. C4‐grass cover was positively correlated with fire frequency and negatively correlated with tree canopy cover. C3‐grass cover was highest at 40% tree canopy cover on unburned sites and at 60% tree canopy cover on frequently burned sites. Total forb cover was maximized at fire frequencies of 4–7 fires per decade, but was not significantly influenced by tree canopy cover. Cover of N‐fixing forbs was highest in shaded areas, particularly on frequently burned sites, while combined cover of all other forbs was negatively correlated with tree canopy cover. Conclusions: The relative influences of fire frequency and tree canopy cover on understorey plant functional group cover vary among plant functional groups, but both play a significant role in structuring savanna and woodland understorey vegetation. When restoring degraded savannas, direct manipulation of overstorey tree canopy cover should be considered to rapidly reduce shading from fire‐resistant overstorey trees. Prescribed fires can then be used to suppress understorey woody plants and promote establishment of light‐demanding grasses and forbs.     

Can plant–natural enemy communication withstand disruption by biotic and abiotic factors?

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Reintroduction of Wild Lupine (Lupinus perennis L.) Depends on Variation in Canopy,Vegetation, and Litter Cover

We experimentally examined the effects of canopy, vegetation, and leaf litter cover on the demography of Wild lupines (Lupinus perennis) in a central North American oak savanna spanning 9 years. We also compared the distribution of Wild lupine across the landscape to results predicted by the demographic experiments. With less canopy cover, soil temperatures were warmer and seedlings emerged earlier. Seedling survival increased 14% with each additional leaf grown. Seedling survival was four times greater in openings and partial shade than in dense shade. Seedling survival was also influenced by interactions between canopy cover and vegetation cover, between canopy cover and leaf litter, and among canopy cover, vegetation cover, and litter cover. In openings, seedlings had higher survival when vegetative cover was present, suggesting a positive shading effect on survival, but with greater canopy cover vegetative cover reduced survival. Seedling survival was greater for plants that experienced herbivory, a result that was probably related to plant size and quality rather than having been eaten. Survival of lupines to 9 years after seed planting was greatest in the partial shade, moderate in openings, and least in dense shade. Wild lupine cover across the landscape was greatest when litter cover was low and canopy cover and ground layer cover were moderate. Reduction of canopy cover by burning or cutting, and reduction of leaf litter by prescribed burning will benefit the reintroduction of Wild lupine by increasing light, reducing litter cover, and creating disturbances; however, the reduction of vegetation cover in openings may hinder lupine reintroduction.     

Volatile terpenes – mediators of plant-to-plant communication

Plants interact with other organisms employing volatile organic compounds (VOCs). The largest group of plant-released VOCs are terpenes, comprised of isoprene, monoterpenes, and sesquiterpenes. Mono- and sesquiterpenes are well-known communication compounds in plant–insect interactions, whereas the smallest, most commonly emitted terpene, isoprene, is rather assigned a function in combating abiotic stresses. Recently, it has become evident that different volatile terpenes also act as plant-to-plant signaling cues. Upon being perceived, specific volatile terpenes can sensitize distinct signaling pathways in receiver plant cells, which in turn trigger plant innate immune responses. This vastly extends the range of action of volatile terpenes, which not only protect plants from various biotic and abiotic stresses, but also convey information about environmental constraints within and between plants. As a result, plant–insect and plant–pathogen interactions, which are believed to influence each other through phytohormone crosstalk, are likely equally sensitive to reciprocal regulation via volatile terpene cues. Here, we review the current knowledge of terpenes as volatile semiochemicals and discuss why and how volatile terpenes make good signaling cues. We discuss how volatile terpenes may be perceived by plants, what are possible downstream signaling events in receiver plants, and how responses to different terpene cues might interact to orchestrate the net plant response to multiple stresses. Finally, we discuss how the signal can be further transmitted to the community level leading to a mutually beneficial community-scale response or distinct signaling with near kin.     

Do biotic interactions modulate ecosystem functioning along stress gradients? Insights from semi-arid plant and biological soil crust communities

Climate change will exacerbate the degree of abiotic stress experienced by semi-arid ecosystems. While abiotic stress profoundly affects biotic interactions, their potential role as modulators of ecosystem responses to climate change is largely unknown. Using plants and biological soil crusts, we tested the relative importance of facilitative–competitive interactions and other community attributes (cover, species richness and species evenness) as drivers of ecosystem functioning along stress gradients in semi-arid Mediterranean ecosystems. Biotic interactions shifted from facilitation to competition along stress gradients driven by water availability and temperature. These changes were, however, dependent on the spatial scale and the community considered. We found little evidence to suggest that biotic interactions are a major direct influence upon indicators of ecosystem functioning (soil respiration, organic carbon, water-holding capacity, compaction and the activity of enzymes related to the carbon, nitrogen and phosphorus cycles) along stress gradients. However, attributes such as cover and species richness showed a direct effect on ecosystem functioning. Our results do not agree with predictions emphasizing that the importance of plant–plant interactions will be increased under climate change in dry environments, and indicate that reductions in the cover of plant and biological soil crust communities will negatively impact ecosystems under future climatic conditions.     

Intraspecific competition replaces interspecific facilitation as abiotic stress decreases: The shifting nature of plant–plant interactions

Plant–plant interactions change depending on environmental conditions, shifting from competition to facilitation when the stress is high. In addition to these changes, the relevance of intraspecific compared to interspecific interactions may also shift as abiotic stress does. We inferred intra- and interspecific plant–plant interactions of the cushion plant Hormathophylla spinosa as related to the dominant shrub Juniperus sabina in two sites with contrasting abiotic conditions (a slope with high-stress conditions vs. a valley bottom with milder conditions) in a Mediterranean high mountain. Specifically, we studied the spatial patterns and several variables related to plant performance (plant size and form, non-structural carbohydrate – NSC – concentrations and radial growth) of H. spinosa.The spatial pattern varied depending on site conditions. H. spinosa plants were positively associated with juniper in the high-stress slope site, probably through higher establishment rates due to the amelioration of soil conditions. In contrast, in the milder valley site H. spinosa establishment occurred mostly in open areas. Age structure, inferred from annual rings, reflected a massive establishment event in the whole study area which occurred 30–50 years ago. Canopy variables and radial growth were density dependent: both were negatively affected by the high density of H. spinosa individuals in the valley, but favoured by junipers on the slope. Interestingly, NSCs showed the opposite pattern, suggesting lower investment in growth by H. spinosa plants in the valley than on the slope.Our results reinforce the strong links existing between intra- and interspecific relationships and the need to include both when studying the influence of abiotic conditions on plant–plant interactions. This approach enabled us to detect that the direction and intensity of plant–plant interactions may shift at different ecological levels. Particularly interesting was the finding that optimal sites at the population level may not necessarily be the sites showing maximum individual performance.     

Birch effects on root fungal colonisation of crowberry are uniform along different environmental gradients

Changes in plant–fungal interactions were often suggested as one of possible mechanisms behind facilitative plant–plant effects in harsh environments. We asked how the mycorrhizal and dark septate endophyte (DSE) colonisations of understorey crowberry (Empetrum nigrum ssp. hermaphroditum) are affected by proximity to mature mountain birch trees (Betula pubescens ssp. czerepanovii) along three abiotic stress gradients (pollution, elevation, seashore) in the Kola peninsula, NW Russia. Stress level affected shoot growth and reproduction in crowberry, but had no effect on root fungal colonisation. In contrast, proximity to a mountain birch tree had no effect on either growth or reproduction of crowberry, but changed all characteristics of root colonisation. The mycorrhizal coil colonisation of crowberry was on average 21% higher near a birch tree, whereas other parameters were higher outside of canopy area (hyaline hyphae: 12%; DSE hyphae: 16%; DSE sclerotia: 42%). Effects of birch tree on root fungal colonisation in crowberry did not depend on the level of abiotic stress. Although we detected a weak positive association between growth of crowberry and its mycorrhizal coil colonisation, we conclude that mycorrhizal and DSE colonisations of crowberry are primarily affected by the abiotic environment. None of the detected patterns was consistent with the patterns expected from the theories concerning stress effects on plant–plant interactions.     

Interspecific facilitation and critical transitions in arid ecosystems

Climate change and intensified land‐use impose severe stress on arid ecosystems, resulting in relatively rapid degradation which is difficult to reverse. To prevent such critical transitions it is crucial to detect early warning signals. Increased ‘patchiness’– smaller and fewer vegetated patches – is thought to be such a signal, but the underlying mechanisms are still poorly understood. Facilitation between plants is known to be an important mechanism driving the patchiness of the vegetation, but we lack understanding of how interactions between plants change in response to combined effects of drought and consumer pressure – the main stressors in many arid ecosystems. Over the last decade numerous experimental studies have tested how intensity of facilitation between plants changes with increasing stress. The most recent synthesis predicts a decline in facilitation intensity at the severe end of a drought stress gradient. Adding consumer pressure may result in even earlier and faster declines in facilitation intensity. So far, studies on critical transitions and plant–plant interactions have developed separately. The relationship between stress and facilitation intensity has been overlooked in critical transition theory, while facilitation intensity may determine the position of a critical transition threshold. In this study, we incorporate experimental studies on the relation between stress and facilitation intensity into the critical transition framework, to improve our ability to predict critical transitions. Moreover, we propose that a decline in facilitation intensity at the severe end of a stress gradient may occur prior to a critical transition. Inclusion of consumer pressure will speed up this process, leading to earlier and faster degradation. In‐field monitoring of seedling–facilitator associations and declines in facilitator recruitment can indicate declines in facilitation intensity and may thus provide additional early warning signals for imminent critical transitions, besides increased patchiness.     

The shrub Rhodomyrtus tomentosa acts as a nurse plant for seedlings differing in shade tolerance in degraded land of South China

Questions: What are the nurse effects of Rhodomyrtus tomentosa in degraded land of South China? Are canopy or soil factors responsible for the main nurse effect? Do facilitative effects increase with the shade tolerance of the target species? Location: Degraded shrubland in South China. Methods: Seedlings of three native climax woody species (Schima superba, Michelia macclurei, Castanopsis fissa) that differ in shade tolerance were subjected to four treatments by transplantation: (1) under the canopy of R. tomentosa shrubs; (2) in open interspaces without vegetation cover (control); (3) under the canopy of R. tomentosa from which canopies had been removed; and (4) in open interspaces without vegetation but covered by branches and leaves of R. tomentosa. Results: At low soil nutrient levels, increased canopy shade, soil porosity and soil moisture under the canopy of R. tomentosa increased seedling survival of the climax tree species S. superba, C. fissa and M. macclurei, and shoot height of S. superba. The nurse effect (a form of facilitation) of R. tomentosa depended more on canopy shade than on soil amelioration. The magnitude of facilitation or nurse effect was positively correlated with shade tolerance of the target species. Conclusions: Use of nurse plants in restoration differs from traditional reforestation (clearing and/or burning to reduce interspecific competition between target tree species and non‐target species) because it focuses on positive interactions between nurse plants and target plants that increase establishment of target species and reduce time required for restoration. Because nurse effects of R. tomentosa shrubs tended to be larger on target species with greater shade tolerance, shade‐tolerant plants are suggested as target species to accelerate restoration.     

Influence of cattle grazing practices on forest understorey structure in north‐eastern New South Wales

Abstract In eastern Australia the practice of grazing cattle in eucalypt forests and woodlands, as a supplementary activity to farmland grazing, is widespread. It is typically accompanied by burning at frequent intervals by graziers to promote more nutritious and digestible growth of the ground cover for their livestock. Collectively, these forest grazing practices affect understorey structure, which in turn affects other biotic and abiotic components of these ecosystems. In order to test how significant the effects of forest grazing practices are relative to the effects of other management practices and environmental variables and the degree to which grazing practices determine understorey vegetation structure, we surveyed 58 sites on the northern tablelands of New South Wales, Australia. All sites were located in eucalypt forest and were stratified by grazing status (presence or absence): time since logging, time since wildfire, geology, aspect, slope and topographic position. At each site an index of vegetation complexity and the most abundant plant species were recorded. The data were analysed by a backwards stepwise multiple regression. Grazing practices had the greatest influence on understorey vegetation complexity of any of the measured attributes. The grazed sites were characterized by a significantly lower vegetation complexity score, different dominant understorey species, reduced or absent shrub layers, and an open, simplified and more grassy understorey structure compared with ungrazed sites. Time since logging and time since wildfire also significantly affected understorey structure. Our results indicate that cattle grazing practices (i.e. grazing and the associated frequent fire regimes) can have major effects on forest structure and composition at a regional level.     

The Art of Being Flexible: How to Escape from Shade,Salt, and Drought

Environmental stresses, such as shading of the shoot, drought, and soil salinity, threaten plant growth, yield, and survival. Plants can alleviate the impact of these stresses through various modes of phenotypic plasticity, such as shade avoidance and halotropism. Here, we review the current state of knowledge regarding the mechanisms that control plant developmental responses to shade, salt, and drought stress. We discuss plant hormones and cellular signaling pathways that control shoot branching and elongation responses to shade and root architecture modulation in response to drought and salinity. Because belowground stresses also result in aboveground changes and vice versa, we then outline how a wider palette of plant phenotypic traits is affected by the individual stresses. Consequently, we argue for a research agenda that integrates multiple plant organs, responses, and stresses. This will generate the scientific understanding needed for future crop improvement programs aiming at crops that can maintain yields under variable and suboptimal conditions.A fundamental difference between plant and animal development is the plasticity in organ formation after germination. Whereas animals are born with a complete set of organs, a germinating seedling has just one embryonic root and one or two embryonic leaves, the cotyledons. All other organs are formed postembryonically, by the interplay of developmental programs and environmental conditions. So, although each plant has a basic body plan, its final size and shape are largely determined by the specific conditions that the plant experiences, and its growth can be adjusted to suit those conditions. This interplay is crucial in both natural and agricultural settings where plants forage for resources and often avoid/escape from stress.Examples of how plants adjust to environmental conditions include phototropism (Darwin, 1880) to bring the photosynthesizing leaves into well-lit microsites such as canopy gaps and root proliferation toward moisture- or nutrient-rich areas to enhance water uptake and nutrient acquisition (Comas et al., 2013). Examples of stress escape include shoot elongation away from the shade of neighbor plants (shade avoidance; Pierik and de Wit, 2014), escape from submerged conditions to reach the air (Bailey-Serres and Voesenek, 2008), and root growth away from saline soil microsites (halotropism; Galvan-Ampudia et al., 2013). Although some of these responses are termed escape from stress (e.g. shade avoidance), others are considered as attraction to more favorable conditions (e.g. hydrotropism). In the case of directional growth responses, the most unifying way is probably to consider these as responses to gradients of stresses (e.g. salt) or resources (e.g. water).The molecular, biochemical, and physiological pathways that underlie these responses have been intensively researched, and this has provided substantial knowledge on the regulatory mechanisms. However, relatively little research has been devoted to studying these modes of plasticity in combination. For example, dense plantings of crops growing on irrigated soils in arid conditions likely need to deal with drought, soil salinity, and shading by neighbor crops and weeds simultaneously. Above ground, plants use light cues, particularly enrichment of far-red light (FR) through reflection by nearby vegetation, to detect neighboring vegetation and respond with shade avoidance responses (Casal, 2013; Pierik and de Wit, 2014). Below ground, plants can sense neighbors and their abiotic environment through a variety of putative cues. Some of these result from selective changes made to the rhizospheres by root absorption of minerals and water and excretion of organic compounds. Plants respond to these cues in various ways, including growth toward or away from neighbors, nutrient hotspots, water, and more (Fang et al., 2013; Pierik et al., 2013).Importantly, the global crop production chain is anticipating intensification of various abiotic stresses: increased temperatures, progressive salinization of highly water-limited production grounds, and more extreme situations of drought and flood (Tubiello et al., 2007; Bailey-Serres and Voesenek, 2008; Munns and Tester, 2008). At the same time, agricultural productivity must be increased to feed the ever-expanding global population, calling for high-density cropping systems with potentially severe mutual shading among plants. Therefore, it is of great importance to understand how plants respond to high-density and abiotic stress(es) simultaneously.Here, we will review the current molecular and physiological understanding of both shoot developmental plasticity in response to high plant density-derived light signals (shade avoidance) and root developmental plasticity in response to the widely occurring abiotic stresses salt and drought. We will then implement this mechanistic knowledge to generate ideas about (1) how these different modes of plasticity may interact to modulate the known stress response phenotypes and (2) how responses to one stress may affect responses to a second. Addressing these ideas experimentally will generate the knowledge needed to guide crop improvement programs under suboptimal agricultural conditions.     

Within‐stand spatial structure and relation of boreal canopy and understorey vegetation

Question: How do spatial patterns and associations of canopy and understorey vegetation vary with spatial scale along a gradient of canopy composition in boreal mixed‐wood forests, from younger Aspen stands dominated by Populus tremuloides and P. balsamifera to older Mixed and Conifer stands dominated by Picea glauca? Do canopy evergreen conifers and broad‐leaved deciduous trees differ in their spatial relationships with understorey vegetation? Location: EMEND experimental site, Alberta, Canada. Methods: Canopy and understorey vegetation were sampled in 28 transects of 100 contiguous 0.5 m × 0.5 m quadrats in three forest stand types. Vegetation spatial patterns and relationships were analysed using wavelets. Results: Boreal mixed‐wood canopy and understorey vegetation are patchily distributed at a range of small spatial scales. The scale of canopy and understorey spatial patterns generally increased with increasing conifer presence in the canopy. Associations between canopy and understorey were highly variable among stand types, transects and spatial scales. Understorey vascular plant cover was generally positively associated with canopy deciduous tree cover and negatively associated with canopy conifer tree cover at spatial scales from 5–15 m. Understorey non‐vascular plant cover and community composition were more variable in their relationships with canopy cover, showing both positive and negative associations at a range of spatial scales. Conclusions: The spatial structure and relation of boreal mixed‐wood canopy and understorey vegetation varied with spatial scale. Differences in understorey spatial structure among stand types were consistent with a nucleation model of patch dynamics during succession in boreal mixed‐wood forests.     

Canopy Openness Enhances Diversity of Ant–Plant Interactions in the Brazilian Amazon Rain Forest

In closed‐canopy tropical forest understory, light availability is a significant determinant of habitat diversity because canopy structure is highly variable in most tropical forests. Consequently, variation in canopy cover affects the composition and distribution of plant species via creating variable light environments. Nevertheless, little is known about how variation in canopy openness structures patterns of plant–animal interactions. Because of the great diversity and dominance of ants in tropical environments, we used ant–plant interactions as a focal network to evaluate how variation in canopy cover influences patterns of plant–insect interactions in the Brazilian Amazon rain forest. We observed that small increases in canopy openness are associated with increased diversity of ant–plant interactions in our study area, and this change is independent of plant or ant species richness. Additionally, we found smaller niche overlap for both ants and plants associated with greater canopy openness. We hypothesize that enhanced light availability increases the breadth of ant foraging sources because variation in light availability gives rise to plant resources of different quality and amounts. Moreover, greater light availability promotes vegetative growth in plants, creating ant foraging ‘bridges’ between plants. In sum, our results highlight the importance of environmental heterogeneity as a determinant of ant–plant interaction diversity in tropical environments.     

Plant–rhizobacteria interactions alleviate abiotic stress conditions

Root-colonizing non-pathogenic bacteria can increase plant resistance to biotic and abiotic stress factors. Bacterial inoculates have been applied as biofertilizers and can increase the effectiveness of phytoremediation. Inoculating plants with non-pathogenic bacteria can provide 'bioprotection' against biotic stresses, and some root-colonizing bacteria increase tolerance against abiotic stresses such as drought, salinity and metal toxicity. Systematic identification of bacterial strains providing cross-protection against multiple stressors would be highly valuable for agricultural production in changing environmental conditions. For bacterial cross-protection to be an effective tool, a better understanding of the underlying morphological, physiological and molecular mechanisms of bacterially mediated stress tolerance, and the phenomenon of cross-protection is critical. Beneficial bacteria-mediated plant gene expression studies under non-stress conditions or during pathogenic rhizobacteria–plant interactions are plentiful, but only few molecular studies on beneficial interactions under abiotic stress situations have been reported. Thus, here we attempt an overview of current knowledge on physiological impacts and modes of action of bacterial mitigation of abiotic stress symptoms in plants. Where available, molecular data will be provided to support physiological or morphological observations. We indicate further research avenues to enable better use of cross-protection capacities of root-colonizing non-pathogenic bacteria in agricultural production systems affected by a changing climate.     

Plant adaptation to dynamically changing environment: the shade avoidance response

The success of competitive interactions between plants determines the chance of survival of individuals and eventually of whole plant species. Shade-tolerant plants have adapted their photosynthesis to function optimally under low-light conditions. These plants are therefore capable of long-term survival under a canopy shade. In contrast, shade-avoiding plants adapt their growth to perceive maximum sunlight and therefore rapidly dominate gaps in a canopy. Daylight contains roughly equal proportions of red and far-red light, but within vegetation that ratio is lowered as a result of red absorption by photosynthetic pigments. This light quality change is perceived through the phytochrome system as an unambiguous signal of the proximity of neighbors resulting in a suite of developmental responses (termed the shade avoidance response) that, when successful, result in the overgrowth of those neighbors. Shoot elongation induced by low red/far-red light may confer high relative fitness in natural dense communities. However, since elongation is often achieved at the expense of leaf and root growth, shade avoidance may lead to reduction in crop plant productivity. Over the past decade, major progresses have been achieved in the understanding of the molecular basis of shade avoidance. However, uncovering the mechanisms underpinning plant response and adaptation to changes in the ratio of red to far-red light is key to design new strategies to precise modulate shade avoidance in time and space without impairing the overall crop ability to compete for light.     

Allocation,stress tolerance and carbon transport in plants: how does phloem physiology affect plant ecology?

Despite the crucial role of carbon transport in whole plant physiology and its impact on plant–environment interactions and ecosystem function, relatively little research has tried to examine how phloem physiology impacts plant ecology. In this review, we highlight several areas of active research where inquiry into phloem physiology has increased our understanding of whole plant function and ecological processes. We consider how xylem–phloem interactions impact plant drought tolerance and reproduction, how phloem transport influences carbon allocation in trees and carbon cycling in ecosystems and how phloem function mediates plant relations with insects, pests, microbes and symbiotes. We argue that in spite of challenges that exist in studying phloem physiology, it is critical that we consider the role of this dynamic vascular system when examining the relationship between plants and their biotic and abiotic environment.     

Are variations of direct and indirect plant interactions along a climatic gradient dependent on species’ strategies? An experiment on tree seedlings

Investigating how interactions among plants depend on environmental conditions is key to understand and predict plant communities’ response to climate change. However, while many studies have shown how direct interactions change along climatic gradients, indirect interactions have received far less attention. In this study, we aim at contributing to a more complete understanding of how biotic interactions are modulated by climatic conditions. We investigated both direct and indirect effects of adult tree canopy and ground vegetation on seedling growth and survival in five tree species in the French Alps. To explore the effect of environmental conditions, the experiment was carried out at 10 sites along a climatic gradient closely related to temperature. While seedling growth was little affected by direct and indirect interactions, seedling survival showed significant patterns across multiple species. Ground vegetation had a strong direct competitive effect on seedling survival under warmer conditions. This effect decreased or shifted to facilitation at lower temperatures. While the confidence intervals were wider for the effect of adult canopy, it displayed the same pattern. The monitoring of micro‐environmental conditions revealed that competition by ground vegetation in warmer sites could be related to reduced water availability; and weak facilitation by adult canopy in colder sites to protection against frost. For a cold‐intolerant and shade‐tolerant species (Fagus sylvatica), adult canopy indirectly facilitated seedling survival by suppressing ground vegetation at high temperature sites. The other more cold tolerant species did not show this indirect effect (Pinus uncinata, Larix decidua and Abies alba). Our results support the widely observed pattern of stronger direct competition in more productive climates. However, for shade tolerant species, the effect of direct competition may be buffered by tree canopies reducing the competition of ground vegetation, resulting in an opposite trend for indirect interactions across the climatic gradient.     

Crop Improvement and Abiotic Stress Tolerance Promoted by Moringa Leaf Extract

Moringa leaf extract (MLE) has been shown to promote beneficial outcomes in animals and plants. It is rich in amino acids, antioxidants, phytohormones, minerals, and many other bioactive compounds with nutritional and growth-promoting potential. Recent reports indicated that MLE improved abiotic stress tolerance in plants. Our understanding of the mechanisms underlying MLE-mediated abiotic stress tolerance remains limited. This review summarizes the existing literature on the role of MLE in promoting plant abiotic stress acclimation processes. MLE is applied to plants in a variety of ways, including foliar spray, rooting media, and seed priming. Exogenous application of MLE promoted crop plant growth, photosynthesis, and yield under both nonstress and abiotic stress conditions. MLE treatment reduced the severity of osmotic and oxidative stress in plants by regulating osmolyte accumulation, antioxidant synthesis, and secondary metabolites. MLE also improves mineral homeostasis in the presence of abiotic stress. Overall, this review describes the potential mechanisms underpinning MLE-mediated stress tolerance.