The ectomycorrhizal community in urban linden trees and its relationship with soil properties

The aim of this study was to verify if root tip vitality and ectomycorrhization in 16 mature urban linden (Tilia vulgaris Hayne) trees, growing in roadside and park side sites, at moderately and strongly declining levels, could be associated with main soil properties. From the 738 soil and root samples collected over the four study periods, 52 ectomycorrhizae were recorded. The study demonstrated that the number of non-vital, vital non-mycorrhizal and mycorrhizal tips never differed significantly, neither among samples collected beneath the same tree nor among trees belonging to the same site and decline class, with the exception of the number of vital non-mycorrhizal tips showing significant differences among sites. Significant differences in ectomycorrhizal community composition between sites and between decline classes were found. This result may be related to the variation of the environmental variables as a whole rather than to the influence of a single or few factors, even if capability, permanent wilting point and available water are the soil parameters most significantly related to diversity in ectomycorrhizal community.     

A geostatistical model to describe root vitality and ectomycorrhization in Norway spruce

Abstract

The spatial distribution of vital root tips and ectomycorrhizal (ECM) communities in forest soils is characterized by patchiness at a microscale level, mostly related to the distribution patterns of biotic and abiotic factors. A geostatistical model was applied to verify if spatial analyses could be useful in identifying an appropriate sampling method to study root tip vitality, ectomycorrhization and the ECM community. Root samples were collected from two high mountain Norway spruce forests (Trentino province, Italy) following a geometrical design. Laboratory microscopic and geostatistical ordinary kriging analyses were used to map tip vitality and ectomycorrhization degree, ECM richness and distribution grouped in “exploration types” (amount of emanating hyphae or presence and differentiation of rhizomorphs). Spatial gradients of the examined features existed at plant level, associated to the up-downslope direction (root tip vitality and ectomycorrhization, ECM richness) and distance from the stem base (ECM exploration types). The effectiveness of the geostatistical model used demonstrates that a geometrical sampling design, associated to spatial mapping techniques, can be useful in research where the tree, and not the forest, is the subject (mycological and phytopathological studies).     

Fine-scale distribution of ectomycorrhizal fungi colonizing Tsuga diversifolia seedlings growing on rocks in a subalpine Abies veitchii forest

Numerous species of ectomycorrhizal (ECM) fungi coexist under the forest floor. To explore the mechanisms of coexistence, we investigated the fine-scale distribution of ECM fungal species colonizing root tips in the root system of Tsuga diversifolia seedlings in a subalpine forest. ECM root tips of three seedlings growing on the flat top surface of rocks were sampled after recording their positions in the root system. After the root tips were grouped by terminal-restriction fragment length polymorphism (T-RFLP) analysis of ITS rDNA, the fungal species representing each T-RFLP group were identified using DNA sequencing. Based on the fungal species identification, the distribution of root tips colonized by each ECM fungus was mapped. Significant clustering of root tips was estimated for each fungal species by comparing actual and randomly simulated distributions. In total, the three seedlings were colonized by 40 ECM fungal species. The composition of colonizing fungal species was quite different among the seedlings. Twelve of the 15 major ECM fungal species clustered significantly within a few centimeters. Some clusters overlapped or intermingled, while others were unique. Areas with high fungal species diversity were also identified in the root system. In this report, the mechanisms underlying generation of these ECM root tip clusters in the root system are discussed.     

Ectomycorrhizal root development in wet Alder carr forests in response to desiccation and eutrophication

Effects of desiccation and eutrophication on ectomycorrhizal (ECM) root development in wet Alder carr forests in The Netherlands were studied. In northwestern Europe, wet Alder carr forests are found mostly in peatlands and along streams, forming an important component of wetland ecosystems. The dominant tree species in wet Alder carr forests is Alnus glutinosa (L.) Gaertn. (Black alder), which associates with ectomycorrhizal fungi. During recent decades, wet Alder carr forests in Europe have declined because of desiccation and eutrophication, particularly in The Netherlands. In the present study, the number of root tips of A. glutinosa trees was highest in an undisturbed wet Alder carr forest in a peatland area. Eutrophication in the peatland area significantly inhibited ectomycorrhizal (ECM) root development of A. glutinosa. In the eutrophied forest, ECM root tips were observed only close to A. glutinosa trees growing on hummocks. The concentrations of nitrate and potassium in soil water of the eutrophied forest were significantly higher than in the undisturbed forest, while magnesium and iron concentrations and the pH were significantly lower. The number of ECM root tips of A. glutinosa in a desiccated forest along a stream was generally lower than in an undisturbed wet Alder carr forest on waterlogged soil in the same area. The sulphate concentration in soil water in the desiccated forest was significantly higher than in the forest on waterlogged soil. ECM root development of A. glutinosa may have been negatively affected by the chemical composition of the soil water.     

Does ectomycorrhizal fungal community structure vary along a Japanese black pine (Pinus thunbergii) to black locust (Robinia pseudoacacia) gradient?

In this study we examined the role of the nitrogen-fixing tree, Robinia pseudoacacia (black locust), in ectomycorrhizal (ECM) formation and ECM community of Pinus thunbergii (Japanese black pine) seedlings. Two 200 m(2) experimental plots were established at the border between a Japanese black pine- and a black locust-dominated area in a coastal forest. The ECM fungal community of pine seedlings was examined by PCR-RFLP and sequence analysis. We analyzed the relationship between ECM formation, ECM community, growth, and nutrient status of pine seedlings and environmental conditions using the Mantel test and structural equation model. Percentages of ECM root tips, the number of ECM fungal species and ECM diversity on pine seedlings decreased in the black locust-dominated area. Cenococcum geophilum and Russula spp. were dominant in the Japanese black pine-dominated area, whereas Tomentella spp. were dominant in the black locust-dominated area. Nitrogen (N) concentration in soils or pine seedlings strongly influenced the percentage of ECM root tips, the number of ECM fungal species and ECM fungal similarity. These results imply the long-term eutrophication caused by N-fixing trees can change ECM formation and ECM community structure.     

Long-term effect of apatite on ectomycorrhizal growth and community structure

Ectomycorrhizal (ECM) fungi are efficient at taking up phosphorus (P) from mineral sources, such as apatite, which are not easily available to the host trees. Since ECM fungal species differ in P uptake rates, it can be expected that the composition of the ECM fungal community will change upon exposure to apatite, provided that the P transfer is rewarded by more carbon being transferred to the fungal symbiont. Control and apatite-amended mesh bags were buried in pairs in the humus layer of a P-poor Norway spruce forest. The ECM fungal community that colonized these bags was analyzed by DNA extraction, PCR amplification of the internal transcribed spacer (ITS) region, cloning, and random sequencing. Fungal biomass was estimated by ergosterol analysis. No change in the ECM fungal community structure was seen after 5?years of apatite exposure, although the fungal biomass increased threefold upon apatite amendment. Our results indicate that host trees enhance carbon allocation to ECM fungi colonizing P sources in P-poor forests but the lack of change in the composition of the ECM fungal community suggests that P transfer rates were similar among the species. Alternatively, higher P transfer among certain species was not rewarded with higher carbon transfer from the host.     

Floristic diversity of an eastern Mediterranean dwarf shrubland: the importance of soil pH

Questions: Does plant species richness and composition of eastern Mediterranean dwarf shrubland (phrygana) correlate with soil pH? How important is the effect of pH on species diversity in relation to other environmental factors in this ecosystem? What is the evolutionary background of the diversity–pH relationship? Location: Western Crete, Greece. Methods: Species composition of vascular plants, soil and other environmental variables were sampled in 100‐m² plots on acidic and basic bedrock in phrygana vegetation. The relationships between species composition and environmental variables (including climate) were tested using canonical correspondence analysis, and relationships between species richness and environment using correlation and regression analyses. Data were analysed separately for different plant functional types based on life form and life span. Results: Although soil pH varied across a narrow range (5.9‐8.1), species composition changed significantly along the pH gradient within all plant functional types. For most functional types, the effect of soil pH on species composition was stronger than that of other environmental variables. Species richness of annuals, geophytes and suffruticose chamaephytes increased with soil pH, while richness of hemicryptophytes and shrubs was not correlated with pH. Conclusions: The results are consistent with the evolutionary species pool hypothesis. High numbers of calcicole annuals, geophytes and suffruticose chamaephytes may be a result of the evolution of these groups on base‐rich dry soils in the Mediterranean climate. In contrast, hemicryptophytes, a life form typical of the temperate zone, evolved on both acidic and basic soils and therefore their species numbers do not respond to soil pH across the narrow range studied. The lack of a relationship between shrub species richness and pH is difficult to explain: it may reflect the more diverse or older origin of Mediterranean woody species and their conservative niches.     

Post-fire, seasonal and annual dynamics of the ectomycorrhizal community in a Quercus ilex L. forest over a 3-year period

Two study plots, burned and control, were established in autumn 1998 in a Quercus ilex forest located in northern Spain, part of which had been affected by a low intensity fire in 1994. Soil samples for ectomycorrhizae (ECM) were taken over a 3-year period in each study plot in spring, summer, autumn and winter. ECM morphotypes were identified and the relative abundance of each morphotype in each soil sample calculated, along with species richness, Shannon diversity index and percentage of mycorrhization in each soil sample. The relative abundance of certain ECM morphotypes differed between burned and control plots, and the percentage of mycorrhizal tips was significantly lower in the burned than in the control plot. Nevertheless, there were no significant differences in the diversity, species richness or species composition of the ECM community in the burned and control plots. The dominant ECM morphotypes in both stands were Cenococcum geophilum and several thelephoroid fungi. Sphaerosporella brunnea and Pisolithus tinctorius thrived especially in the burned plot, whereas three ectomycorrhizal morphotypes assigned to the genus Hebeloma were especially abundant in the control plot. There was no significant variation in the relative abundance of the ECM morphotypes between seasons, but ECM community species richness was highest in autumn and lowest in summer. The percentage of mycorrhizal tips reached a maximum in winter, with its minimum in autumn. Collection of samples over the 3-year period also enabled us to detect a significant increase in percentage of ECM colonisation in the burned stand over time.     

Fungal diversity in ectomycorrhizal communities: sampling effort and species detection

A number of recent review articles on ectomycorrhizal (ECM) fungal community diversity have highlighted the unprecedented increase in the number of publications on this ecologically important but neglected area. The general features of these species-rich, highly dynamic and complex communities have been comprehensively covered but one aspect crucial to our assessment of diversity, namely the sampling of ECM communities has received less attention. This is a complex issue with two principal components, the physical sampling strategy employed and the life cycle traits of the ECM fungi being examined. Combined, these two components provide the image that we perceive as ECM diversity. This contribution will focus primarily on the former of these components using a recent study from a pine forest in central Sweden to highlight some sampling problems and also to discuss some features common to ECM communities. The two commonly used elements of diversity, species richness and community evenness, present rather different problems in the assessment of ECM diversity. The applicability of using current measures of abundance (number or percentage of root tips colonised) to determine community evenness is discussed in relation to our lack of knowledge on the size of individual genets of ECM fungi. The inherent structure of most ECM communities, with a few common species and a large number of rare species, severely limits our ability to accurately assess species richness. A discussion of theoretical detection limits is included that demonstrates the importance of the sampling effort (no. of samples or tips) involved in assessing species richness. Species area abundance plots are also discussed in this context. It is suggested that sampling strategy (bulk samples versus multiple collections of single tips) may have important consequences when sampling from communities where root tip densities differ. Finally, the need for studies of the spatial distribution of ECM on roots in relation to small-scale soil heterogeneity and of the temporal aspects of ECM community dynamics is raised.     

桂北典型岩溶区植物alpha和beta多样性特征及影响因素

母岩对景观演化和生物定植起着决定性作用,但母岩对区植物多样性是否产生影响是地学、植物学和生态学共同关注的热点。以桂北冠岩、恭城两个典型岩溶区3种母岩(灰岩、碎屑岩、灰岩夹白云岩)天然次生林为研究对象,基于野外调查数据,分析了岩溶区植物alpha和beta多样性特征及影响因素。结果显示:(1)研究区内物种相对丰富,共记录乔木86种、灌木93种、草本56种,灰岩植物alpha多样性最高,碎屑岩次之,灰岩夹白云岩最低;(2)不同母岩和同一母岩区的植物群落相异性均较高,物种周转组分对beta多样性贡献(82.86%-84.49%)显著高于物种丰富度组分(15.51%-17.14%);(3)Pearson和Mantel分析表明,母岩、基岩裸露率和土壤厚度与桂北岩溶区植物alpha和beta多样性显著相关(P<0.05)。综上所述,桂北岩溶森林林分异质性较高,碳酸岩混合沉积区植物多样性显著低于纯灰岩和碎屑岩。植物组成差异主要取决于物种周转过程,母岩、基岩裸露和土壤厚度是主要影响因素。本研究从母岩的角度评估了岩溶森林的植物多样性特征及主要影响因素,为桂北乃至中国西南岩溶地区植被保护和重建提供科学依据和理论支持。     

Changes in ectomycorrhizal fungal community composition and declining diversity along a 2‐million‐year soil chronosequence

Ectomycorrhizal (ECM) fungal communities covary with host plant communities along soil fertility gradients, yet it is unclear whether this reflects changes in host composition, fungal edaphic specialization or priority effects during fungal community establishment. We grew two co‐occurring ECM plant species (to control for host identity) in soils collected along a 2‐million‐year chronosequence representing a strong soil fertility gradient and used soil manipulations to disentangle the effects of edaphic properties from those due to fungal inoculum. Ectomycorrhizal fungal community composition changed and richness declined with increasing soil age; these changes were linked to pedogenesis‐driven shifts in edaphic properties, particularly pH and resin‐exchangeable and organic phosphorus. However, when differences in inoculum potential or soil abiotic properties among soil ages were removed while host identity was held constant, differences in ECM fungal communities and richness among chronosequence stages disappeared. Our results show that ECM fungal communities strongly vary during long‐term ecosystem development, even within the same hosts. However, these changes could not be attributed to short‐term fungal edaphic specialization or differences in fungal inoculum (i.e. density and composition) alone. Rather, they must reflect longer‐term ecosystem‐level feedback between soil, vegetation and ECM fungi during pedogenesis.     

Effects of Liming on Potential Oxalate Secretion and Iron Chelation of Beech Ectomycorrhizal Root Tips

Liming is used to counteract forest decline induced by soil acidification. It consists of Ca and Mg input to forest soil and not only restores tree mineral nutrition but also modifies the availability of nutrients in soil. Ectomycorrhizal (ECM) fungi are involved in mineral nutrient uptake by trees and can recover them through dissolution of mineral surface. Oxalate and siderophore secretion are considered as the main agents of mineral weathering by ECMs. Here, we studied the effects of liming on the potential oxalate secretion and iron complexation by individual beech ECM root tips. Results show that freshly excised Lactarius subdulcis root tips from limed plots presented a high potential oxalate exudation of 177 μM tip⁻¹ h⁻¹. As this ECM species distribution is very dense, it is likely that, in the field, oxalate concentrations in the vicinity of its clusters could be very high. This points out that not only extraradical mycelium but also ECM root tips of certain species can contribute significantly to mineral weathering. Nonmetric multidimensional scaling (NMDS) separated potential oxalate production by ECM root tips in limed and untreated plots, and this activity was mainly driven by L. subdulcis ECMs, but NMDS on potential activity of iron mobilization by ECM root tips did not show a difference between limed and untreated plots. As the mean oxalate secretion did not significantly correlated with the mean iron mobilization by ECM morphotype, we conclude that iron complexation was due to either other organic acids or to siderophores.     

Exploring the homogeneity of terrestrial subterranean communities at a local spatial scale

1. Although caves are generally perceived as isolated habitats, at the local scale, they are often interconnected via a network of fissures in the bedrock. Accordingly, caves in close proximity are expected to host the same, or very similar, animal communities. 2. We explored the extent to which subterranean arthropod communities are homogenous at a local spatial scale of less than 1 km², along with which cave-specific environmental features result in a departure from the expected homogeneous pattern. We approached this question by studying richness and turnover in terrestrial invertebrate communities of 27 caves in a small karst massif in the Western Italian Alps. 3. Specialised subterranean species were homogeneously distributed among caves and were not influenced by seasonality. The only factor driving their distribution was the distance from the cave entrance, with deeper caves yielding a greater diversity of species. 4. Significant spatio-temporal turnover in species not specialised to subterranean life was observed. In summer, there was a significant homogenisation of the community and a more even distribution of species among sites; in winter, these species were missing or found exclusively at greater depths, where environmental conditions were more stable. Furthermore, caves at lower elevations yielded, on average, a greater diversity and abundance of these species. 5. This study demonstrated that the theoretical expectation of no turnover in community composition in caves in close proximity is not always met. Turnover can be mostly attributed to seasonal patterns and sampling depth; thus, our findings have implications for planning sampling and monitoring activities in caves.     

A molecular survey of ectomycorrhizal hyphae in a California Quercus–Pinus woodland

Ectomycorrhizal (ECM) hyphal communities have not been well characterized. Furthermore, there have been few studies where the ECM hyphal community is compared to fungi detected as sporocarps or ECM-colonized root tips. We investigated fungi present as hyphae in a well-studied California Quercus–Pinus woodland. Hyphal species present were compared to those found as sporocarps and ECM root tips at the same site. Hyphae were extracted from root-restrictive nylon mesh in-growth bags buried in the soil near mature Quercus douglasii, Quercus wislizeni, and Pinus sabiniana. Taxa were identified using PCR, cloning, and DNA sequencing of internal transcribed spacer and 28s rDNA. Among the 33 species detected, rhizomorph-forming ECM fungi dominated the hyphal community, especially species of Thelephoraceae and Boletales. Most fungi in soils near Quercus spp. and P. sabiniana were ECM basidiomycetes, but we detected two ECM ascomycetes and three non-mycorrhizal fungi. Many ECM species present as hyphae were also previously detected at this site as sporocarps (18%) or on ECM root tips (58%). However, the hyphal community was mostly dominated by different taxa than either the sporocarp or ECM root communities.     

Changes in ectomycorrhizal community structure on two containerized oak hosts across an experimental hydrologic gradient

Shifts in ectomycorrhizal (ECM) community structure were examined across an experimental hydrologic gradient on containerized seedlings of two oak species, Quercus montana and Quercus palustris, inoculated from a homogenate of roots from mature oak trees. At the end of one growing season, seedlings were harvested, roots were sorted by morphotype, and proportional colonization of each type was determined. DNA was subsequently extracted from individual root tips for polymerase chain reaction, restriction fragment length polymorphism, and rDNA sequencing of the ITS1/5.8S/ITS2 region to determine identities of fungal morphotypes. Twelve distinct molecular types were identified. Analysis of similarity showed that ECM fungal assemblages shifted significantly in composition across the soil moisture gradient. Taxa within the genus Tuber and the family Thelephoraceae were largely responsible for the changes in fungal assemblages. There were also significant differences in ECM community assemblages between the two oak host species. These results demonstrate that the structure of ECM fungal communities depends on both the abiotic and biotic environments and can shift with changes in soil moisture as well as host plant, even within the same genus.     

Nutrient trade‐offs mediated by ectomycorrhizal strategies in plants: Evidence from an Abies species in subalpine forests

1. Ectomycorrhizal (ECM) symbiosis is an evolutionary biological trait of higher plants for effective nutrient uptakes. However, little is known that how the formation and morphological differentiations of ECM roots mediate the nutrients of below‐ and aboveground plant tissues and the balance among nutrient elements across environmental gradients. Here, we investigated the effects of ECM foraging strategies on root and foliar N and P concentrations and N:P ratio Abies faxoniana under variations of climate and soil conditions.
2. The ECM symbionts preferentially mediated P uptake under both N and P limitations. The uptake efficiency of N and P was primarily associated with the ECM root traits, for example, ECM root tip density, superficial area of ECM root tips, and the ratio of living to dead root tips, and was affected by the ECM proliferations and morphological differentiations. The tissue N and P concentrations were positively associated with the abundance of the contact exploration type and negatively with that of the short‐distance exploration type.
3. Our findings indicate that the nutritional status of both below‐ and aboveground plant tissues can be strongly affected by ECM symbiosis in natural environments. Variations in the ECM strategies in response to varying environmental conditions significantly influence plant nutrient uptakes and trade‐offs.

     

Ectomycorrhizal fungal diversity decreases in Mediterranean pine forests adapted to recurrent fires

Fire is a major disturbance linked to the evolutionary history and climate of Mediterranean ecosystems, where the vegetation has evolved fire‐adaptive traits (e.g., serotiny in pines). In Mediterranean forests, mutualistic feedbacks between trees and ectomycorrhizal (ECM) fungi, essential for ecosystem dynamics, might be shaped by recurrent fires. We tested how the structure and function of ECM fungal communities of Pinus pinaster and Pinus halepensis vary among populations subjected to high and low fire recurrence in Mediterranean ecosystems, and analysed the relative contribution of environmental (climate, soil properties) and tree‐mediated (serotiny) factors. For both pines, local and regional ECM fungal diversity were lower in areas of high than low fire recurrence, although certain fungal species were favoured in the former. A general decline of ECM root‐tip enzymatic activity for P. pinaster was associated with high fire recurrence, but not for P. halepensis. Fire recurrence and fire‐related factors such as climate, soil properties or tree phenotype explained these results. In addition to the main influence of climate, the tree fire‐adaptive trait serotiny recovered a great portion of the variation in structure and function of ECM fungal communities associated with fire recurrence. Edaphic conditions (especially pH, tightly linked to bedrock type) were an important driver shaping ECM fungal communities, but mainly at the local scale and probably independently of the fire recurrence. Our results show that ECM fungal community shifts are associated with fire recurrence in fire‐prone dry Mediterranean forests, and reveal complex feedbacks among trees, mutualistic fungi and the surrounding environment in these ecosystems.     

Growth response of drought-stressed Pinus sylvestris seedlings to single- and multi-species inoculation with ectomycorrhizal fungi

Many trees species form symbiotic associations with ectomycorrhizal (ECM) fungi, which improve nutrient and water acquisition of their host. Until now it is unclear whether the species richness of ECM fungi is beneficial for tree seedling performance, be it during moist conditions or drought. We performed a pot experiment using Pinus sylvestris seedlings inoculated with four selected ECM fungi (Cenococcum geophilum, Paxillus involutus, Rhizopogon roseolus and Suillus granulatus) to investigate (i) whether these four ECM fungi, in monoculture or in species mixtures, affect growth of P. sylvestris seedlings, and (ii) whether this effect can be attributed to species number per se or to species identity. Two different watering regimes (moist vs. dry) were applied to examine the context-dependency of the results. Additionally, we assessed the activity of eight extracellular enzymes in the root tips. Shoot growth was enhanced in the presence of S. granulatus, but not by any other ECM fungal species. The positive effect of S. granulatus on shoot growth was more pronounced under moist (threefold increase) than under dry conditions (twofold increase), indicating that the investigated ECM fungi did not provide additional support during drought stress. The activity of secreted extracellular enzymes was higher in S. granulatus than in any other species. In conclusion, our findings suggest that ECM fungal species composition may affect seedling performance in terms of aboveground biomass.     

The functioning of ectomycorrhizal fungi in the field: linkages in space and time

Individual trees, either of the same or different species, can be linked spatially and temporally by the hyphae of ectomycorrhizal (ECM) fungi that allow carbon and nutrients to pass among them and promote forest establishment following disturbance. Spatial and temporal linkages between plants influence the function of ECM fungi in the field. Studies indicate that ECM linkages can reduce plant competition for resources, promote forest recovery, and influence the pattern of plant succession. The degree of influence depends on many factors, including the composition and arrangement of the vegetative community and soil and climatic conditions. Management practices that create intense disturbance and loss of organic matter or promote the introduction of non-ectomycorrhizal host species can decrease the ability of plants to form linkages with ECM fungi. Management practices that retain living trees and shrubs and input of organic matter provide the energy source and substrate necessary for ECM linkages. More research is needed to determine the degree to which ECM fungal linkages occur in the field and their role in ecosystem function and long-term health.     

Assessing the use of sediment organic,carbonate and biogenic silica content as indicators of environmental conditions in Arctic lakes

The objective of this paper is to examine three sediment parameters used in paleolimnological studies, sediment organic, carbonate and biogenic silica (BSi) content, and to quantify the relationship among these parameters and modern environmental conditions for a series of Canadian Arctic lakes. Sediment samples from 63 lakes were analysed for organic, carbonate and BSi content. The environmental and physical parameters of these lakes were recorded. Bedrock composition exerted a very strong influence on sediment organic, BSi and carbonate content, and differences in lake production accounted for much of the remaining variance. The strongest relationships existed among sediment carbonate content, BSi content and water pH. A statistical model developed for predicting water pH provides a new tool for reconstructing past environmental conditions for Arctic aquatic ecosystems. A positive relationship between mean July air temperature and sediment organic and BSi content was detected, but was confounded by effects of bedrock on these parameters. However, the relationship between mean July air temperature and sediment organic and BSi could be even more significant in the context of constant bedrock composition, as is the case in paleolimnological studies.