Although the relationship between species diversity and biomass productivity has been extensively studied in grasslands, the impact of tree species diversity on forest productivity, as well as the ...main drivers of this relationship, are still under discussion. It is widely accepted that the magnitude of the relationship between tree diversity and forest stand productivity is context specific and depends on environmental conditions, but the underlying mechanisms of this relationship are still not fully understood. Competition reduction and facilitation have been identified as key mechanisms driving the diversity–productivity relationship. However, contrasting results have been reported with respect to the extent to which competition reduction and facilitation determine the diversity–productivity relationship. They appear to depend on regional climate, soil fertility, functional diversity of the tree species involved, and developmental stage of the forest. The purpose of this review is to summarize current knowledge and to suggest a conceptual framework to explain the various processes leading to higher productivity of species-rich forests compared with average yields of their respective monocultures. This framework provides three pathways for possible development of the diversity–productivity relationship under a changing climate.
•High structural heterogeneity reduces the diurnal temperature range (DTR).•Effects of stand structure on DTR are more pronounced in regions with low precipitation.•DTR is lower in unmanaged than in ...managed uneven-aged and even-aged stands.•Terrestrial laser scanning based approaches explain 79.4% of variance in DTR.
The microclimate in forest ecosystems can be altered by modifications of stand structure due to forest management or natural forest development. Current forest management practices in Central Europe and North America aim to promote structural heterogeneity and maintain forest canopy cover, which is known to be a major driver of forest microclimate. Here, we investigated the impacts of forest management and structural heterogeneity on the diurnal temperature range (DTR) in 128 managed forest stands in three climatically different locations (Swabian Alb, Hainich-Dün and Schorfheide-Chorin) in Central Europe. Increasing structural heterogeneity by promoting tree size diversity and differentiation increased vertical stratification and resulted in an impaired DTR during the vegetation period. Linear regression models with geographic location, elevation above sea level, canopy openness and measures of structural heterogeneity as explanatory variables explained 79.4–80.9% of variance in DTR. However, the overall effect of structural heterogeneity on DTR was small. Differences in DTR between plots of different main tree species could be attributed to differences in canopy openness and light transmission, whereas tree species diversity had no significant effect on DTR. Unmanaged forests were characterized by a significantly lower DTR than managed, even-aged forests. DTR in uneven-aged stands managed under single tree selection was comparable to unmanaged stands. Terrestrial laser scanning (TLS) derived measures of canopy openness and vertical structure allowed to explain 79.4% of variance in DTR considering geographic location and elevation, which can also be assessed by TLS with integrated GPS and an altimeter. We conclude that structural characteristics of forest stands other than canopy openness contribute marginally to variation in forest microclimate. However, the analyses of structure-microclimate analyses indicate that effects of stand structure on DTR might be more pronounced in regions with low precipitation during the vegetation period.
Quercus robur L. (pedunculate oak) and Quercus petraea (Matt.) Liebl. (sessile oak) are two European oak species of great economic and ecological importance. Even though both oaks have wide ...ecological amplitudes of suitable growing conditions, forests dominated by oaks often fail to regenerate naturally. The regeneration performance of both oak species is assumed to be subject to a variety of variables that interact with one another in complex ways. The novel approach of this research was to study the effect of many ecological variables on the regeneration performance of both oak species together and identify key variables and interactions for different development stages of the oak regeneration on a large scale in the field. For this purpose, overstory and regeneration inventories were conducted in oak dominated forests throughout southern Germany and paired with data on browsing, soil, and light availability. The study was able to verify the assumption that the occurrence of oak regeneration depends on a set of variables and their interactions. Specifically, combinations of site and stand specific variables such as light availability, soil pH and iron content on the one hand, and basal area and species composition of the overstory on the other hand. Also browsing pressure was related to oak abundance. The results also show that the importance of variables and their combinations differs among the development stages of the regeneration. Light availability becomes more important during later development stages, whereas the number of oaks in the overstory is important during early development stages. We conclude that successful natural oak regeneration is more likely to be achieved on sites with lower fertility and requires constantly controlling overstory density. Initially sufficient mature oaks in the overstory should be ensured. In later stages, overstory density should be reduced continuously to meet the increasing light demand of oak seedlings and saplings.
•High tree structural complexity is beneficial to high stand structural complexity.•Most tested measures we observed showed scale dependency.•We found no patterns of emergence for structural ...complexity in forests.•Structural complexity plateaus already with 20 trees grouped together.
Management for complexity has become an important paradigm for European and North American forestry. Recent advancements in data processing allow for a detailed, three-dimensional and objective quantification of structural complexity in forests based on terrestrial laser scanning data. In our study, we used such 3D data from an exemplary temperate broad-leaved forest in Thuringia, Germany, to gain insights to the relationship between tree-level structural complexity and stand-level structural complexity. From our study site, which was 80 by 80 m in extent with a total of 215 trees growing in it, we created a dataset that contained each tree as an independent point cloud. Random sample plots of varying size (10 × 10 m; 15 × 15 m; 20 × 20 m) where used to create sub-plots (sampling with replacement) and to enable for the investigation of effects of scale. Our study revealed that plot-level complexity of plots up to 20 × 20 m is largely determined by the complexity of the most complex-structured tree individual. Furthermore, a high tree complexity and variability thereof in the stand was generally beneficial to stand structural complexity. Other individual tree characteristics, such as a large crowns, were also identified to have positive effects on plot-level complexity. We conclude that management for complexity should focus on large-crowned, highly-complex tree individuals as key elements of stand structural complexity. This indicates that large and old trees may not only be of great importance as habitat trees potentially increasing biodiversity, but also due to their contribution to the overall stand-level complexity.
The complexity of forest structures plays a crucial role in regulating forest ecosystem functions and strongly influences biodiversity. Yet, knowledge of the global patterns and determinants of ...forest structural complexity remains scarce. Using a stand structural complexity index based on terrestrial laser scanning, we quantify the structural complexity of boreal, temperate, subtropical and tropical primary forests. We find that the global variation of forest structural complexity is largely explained by annual precipitation and precipitation seasonality (R² = 0.89). Using the structural complexity of primary forests as benchmark, we model the potential structural complexity across biomes and present a global map of the potential structural complexity of the earth´s forest ecoregions. Our analyses reveal distinct latitudinal patterns of forest structure and show that hotspots of high structural complexity coincide with hotspots of plant diversity. Considering the mechanistic underpinnings of forest structural complexity, our results suggest spatially contrasting changes of forest structure with climate change within and across biomes.
In the past 30 years, many stand structural attributes (SSAs) have been suggested and structural indices have been developed to describe the complex structure of forests. Most studies, however, have ...explored the potential and limits of structural measures to quantify forest structures by applying multiple measures to one stand or few measures to several stands. However, the interdependencies of multiple structural attributes across many stands of different forest management types and developmental stages have not yet been explored. Using 20 structural attributes and 124 completely inventoried 1ha sized stands we tested to what extent structural characteristics reflect different stand types and management intensities, and how these characteristics change over time. We found that single SSAs do not show the clear gradients that they were intended to reflect, suggesting that stand structure should be described by multiple structural attributes, and that these should represent different structural aspects (including vertical, and horizontal heterogeneity, density, and diversity). A principal component analysis showed that combining several SSAs, allowed us to distinguish between stand types. The structure of mature stands remained rather constant over the observed period of about 6 years, while that of young stands changed more rapidly due to ingrowth and mortality. The older the stands, the less the large trees contributed to stand growth relative to their size. We conclude that multiple stand structural attributes are needed to characterise stand types, management effects and to explain stand productivity.
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•A strong effect of site conditions on all fine root variables.•A strong effect of species identity and interaction between site conditions and species identity on fine root ...growth.•No belowground overyielding, but mixing effect observed at specific soil depths.
It is already well known that the effects of tree diversity on aboveground wood productivity depend on tree species identity and site conditions and thus can vary among different tree mixtures and forest sites. The effects of species diversity, specifically on belowground productivity and dynamics, have rarely been studied, so potential interactions with species identity and/or site conditions are largely unknown.
Sequential soil coring was applied in mature pure and mixed stands of European beech, Norway spruce, and Douglas-fir in two regions of northern Germany to investigate the potential site, species, and mixing effects on fine root productivity, mortality, and turnover rates. In these ecologically and economically important species mixtures, a strong effect of site conditions on all fine root variables was found. In particular, more fine root growth was observed at the less favorable site conditions than at the more favorable ones. Species identity and interaction between site conditions and species identity were additional explanatory factors, whereat beech showed the strongest effect by site conditions. No overyielding was observed when fine root productivity was plotted against area potentially available (APA), nor were mortality or turnover. However, at specific soil depths, a mixing effect, caused mainly by beech, was observed for all variables and for both species mixtures.
This study suggests that site conditions and species identity rather than species mixture are essential in explaining fine root dynamics and that increasing tree species diversity may not guarantee higher belowground productivity. However, in the face of climate change and the demand for wood as a natural and renewable resource, an admixture of Douglas-fir to pure European beech stands seems to be a reasonable alternative to an admixture of spruce, a species that is already seriously suffering by drought and bark beetle attacks.
•Literature review on correlations between forest structural attributes and species richness.•85 studies and 410 correlations were analyzed.•Structures related to old-growth successional stage most ...important for many species groups.
The effects of forest management on species richness and diversity have become important research interests. The need to maintain biodiversity for forest ecosystem functioning has led to the question of how strongly and in what ways forest management modifies the diversity and abundance of different species groups. It is well known that many forest species rely on specific structures that may be modified by forest management. Assessing the impact of forest management on species richness may therefore require identification of structural properties. For this literature review we identified a large set of structural attributes that can serve as potential drivers of the richness of different species groups. Most studies included here focused on only one or a few structural attributes as explanatory variables and a limited number of species groups as dependent variables; we therefore analyzed the available publications across species and structural properties. We gathered 410 relationships of structure and species richness out of 85 studies from the temperate region in Europe. Positive, negative, and neutral (non-existent) correlations between species richness and the presence of specific structural properties in European temperate forests were then compiled. Canopy gaps and structural attributes related to old-growth successional stage such as stand age and the share of large old trees were mostly positively correlated with species richness of the different taxa. Especially old-growth structures were ranked high in the reviewed literature. The structural attributes that were mainly positively correlated with species richness or the richness of groups of species may be used for further development of biodiversity monitoring concepts and forest management.
Extensive analyses of soluble primary and secondary metabolites as well as cell wall-bound phenolics of
Medicago truncatula roots by GC–MS, HPLC, and LC–MS during the symbiotic interaction with the ...arbuscular mycorrhizal fungus
Glomus intraradices are reported. Metabolite data sets were subjected to bioinformatics approaches including HCA, PCA and network analyses.
Metabolite profiling of soluble primary and secondary metabolites, as well as cell wall-bound phenolic compounds from roots of barrel medic (
Medicago truncatula) was carried out by GC–MS, HPLC and LC–MS. These analyses revealed a number of metabolic characteristics over 56 days of symbiotic interaction with the arbuscular mycorrhizal (AM) fungus
Glomus intraradices, when compared to the controls, i.e. nonmycorrhizal roots supplied with low and high amounts of phosphate. During the most active stages of overall root mycorrhization, elevated levels of certain amino acids (Glu, Asp, Asn) were observed accompanied by increases in amounts of some fatty acids (palmitic and oleic acids), indicating a mycorrhiza-specific activation of plastidial metabolism. In addition, some accumulating fungus-specific fatty acids (palmitvaccenic and vaccenic acids) were assigned that may be used as markers of fungal root colonization. Stimulation of the biosynthesis of some constitutive isoflavonoids (daidzein, ononin and malonylononin) occurred, however, only at late stages of root mycorrhization. Increase of the levels of saponins correlated AM-independently with plant growth. Only in AM roots was the accumulation of apocarotenoids (cyclohexenone and mycorradicin derivatives) observed. The structures of the unknown cyclohexenone derivatives were identified by spectroscopic methods as glucosides of blumenol C and 13-hydroxyblumenol C and their corresponding malonyl conjugates. During mycorrhization, the levels of typical cell wall-bound phenolics (e.g. 4-hydroxybenzaldehyde, vanillin, ferulic acid) did not change; however, high amounts of cell wall-bound tyrosol were exclusively detected in AM roots.
Principal component analyses of nonpolar primary and secondary metabolites clearly separated AM roots from those of the controls, which was confirmed by an hierarchical cluster analysis. Circular networks of primary nonpolar metabolites showed stronger and more frequent correlations between metabolites in the mycorrhizal roots. The same trend, but to a lesser extent, was observed in nonmycorrhizal roots supplied with high amounts of phosphate. These results indicate a tighter control of primary metabolism in AM roots compared to control plants. Network correlation analyses revealed distinct clusters of amino acids and sugars/aliphatic acids with strong metabolic correlations among one another in all plants analyzed; however, mycorrhizal symbiosis reduced the cluster separation and enlarged the sugar cluster size. The amino acid clusters represent groups of metabolites with strong correlations among one another (cliques) that are differently composed in mycorrhizal and nonmycorrhizal roots. In conclusion, the present work shows for the first time that there are clear differences in development- and symbiosis-dependent primary and secondary metabolism of
M. truncatula roots.
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