Heterogeneous Light Conditions Research Articles

Genotype‐dependent Strategies to “Overcome” Excessive Light: Insights into Non‐Photochemical Quenching under High Light Intensity

AbstractHigh light (HL) intensities have a significant impact on energy flux and distribution within photosynthetic apparatus. To understand the effect of high light intensity (HL) on the HL tolerance mechanisms in tomatoes, we examined the response of the photosynthesis apparatus of 12 tomato genotypes to HL. A reduced electron transfer per reaction center (ET0/RC), an increased energy dissipation (DI0/RC) and non‐photochemical quenching (NPQ), along with a reduced maximum quantum yield of photosystem II (FV/FM), and performance index per absorbed photon (PIABS) were common HL‐induced responses among genotypes; however, the magnitude of those responses was highly genotype‐dependent. Tolerant and sensitive genotypes were distinguished based on chlorophyll fluorescence and energy‐quenching responses to HL. Tolerant genotypes alleviated excess light through energy‐dependent quenching (qE), resulting in smaller photoinhibitory quenching (qI) compared to sensitive genotypes. Quantum yield components also shifted under HL, favoring the quantum yield of NPQ (ՓNPQ) and the quantum yield of basal energy loss (ՓN0), while reducing the efficient quantum yield of PSII (ՓPSII). The impact of HL on tolerant genotypes was less pronounced. While the energy partitioning ratio did not differ significantly between sensitive and tolerant genotypes, the ratio of NPQ components, especially qI, affected plant resilience against HL. These findings provide insights into different patterns of HL‐induced NPQ components in tolerant and sensitive genotypes, aiding the development of resilient crops for heterogeneous light conditions.

Contrasting effects of experiencing temporally heterogeneous light availability versus homogenous shading on plant subsequent responses to light conditions

Temporally heterogeneous environments is hypothesized to correlate with greater plasticity of plants, which has rarely been supported by direct evidence. To address this issue, we subjected three species from different ranges of habitats to a first round of alternating full light and heavy shading (temporally heterogeneous light experience), constant moderate shading and full light conditions (temporally homogeneous light experiences, control) and a second round of light-gradient treatments. We measured plant performance in a series of morphological, biomass, physiological and biochemical traits at the end of each round. Compared to constant full light experience, temporally heterogeneous light conditions induced immediate active biochemical responses (in the first round) with improved late growth in biomass (during the second round); constant moderate shading experience increased photosynthetic physiological and biomass performances of plants in early response, and decreased their late growth in biomass. The karst endemic species of Kmeria septentrionalis showed greater improvement in late growth of biomass and lower decrease in biochemical performance, due to early heterogeneous experience, compared to the non-karst species of Lithocarpus glaber and the karst adaptable species of Celtis sinensis. Results suggested plants will prefer to produce morphological and physiological responses that are less reversible and more costly in the face of more reliable environmental cues at early stage in spite of decreased future growth potential, but to produce immediate biochemical responses for higher late growth potential when early environmental cues are less reliable, to avoid the loss of high costs and low profits. Typical karst species should be more able to benefit from early temporally heterogeneous experience, due to long-term adaptation to karst habitats of high environmental heterogeneity and low resource availability.

Surface light modulation by sea ice and phytoplankton survival in a convective flow model

Plankton dynamics depend in a complex manner on a variety of physical phenomena, according to both experimental and numerical data. In particular, experimental field studies have highlighted the relation between phytoplankton survival and turbulent upwelling and downwelling from thermal convection. Recent numerical works have also shown the importance of accounting for advective transport by persistent structures in simulation models. In nutrient-rich polar marine environments phytoplankton blooms are critically limited by light availability under ice-covered waters. Such heterogeneity of the light intensity distribution, in association with a large-scale coherent fluid flow, can give rise to nontrivial growth dynamics. In this work we extend a previous advection-reaction-diffusion model of phytoplankton light-limited vertical dynamics in the presence of convective transport. Specifically, we consider horizontally heterogeneous light conditions through the use of two regions with different production regimes, modelling the absence (presence) of light under (in between) obstacles. Such a model is intended as an idealized representation of nonuniformly ice-covered polar waters. By means of numerical simulations, we find that the main role of advective transport is to hinder phytoplankton growth, but also that such effect depends on the positions of the obstacles with respect to the upwelling and downwelling flow regions. Furthermore, we show that the sinking speed due to the density difference between phytoplankton organisms and water, while small, plays an important role, which depends on how it adds to the flow. These results indicate that advective transport can have a crucial impact on the survival conditions of sinking phytoplankton species in polar environments.

Parental UV-B radiation regulates the habitat selection of clonal Duchesnea indica in heterogeneous light environments.

Habitat selection behaviour is an effective strategy adopted by clonal plants in heterogeneous understorey light environments, and it is likely regulated by the parental environment's ultraviolet-B radiation levels (UV-B) due to the photomorphogenesis of UV-B and maternal effects. Here, parental ramets of Duchesnea indica were treated with two UV-B radiation levels [high (UV5 group) and low (UV10 group)], newborn offspring were grown under a heterogeneous light environment (ambient light vs shade habitat), and the growth and DNA methylation variations of parents and offspring were analysed. The results showed that parental UV-B affected not only the growth of the parent but also the offspring. The offspring of different UV-B-radiated parents showed different performances. Although these offspring all displayed a tendency to escape from light environments, such as entering shade habitats earlier, and allocating more biomass under shade (33.06% of control, 42.28% of UV5 and 72.73% of UV10), these were particularly obvious in offspring of the high UV-B parent. Improvements in epigenetic diversity (4.77 of control vs 4.83 of UV10) and total DNA methylation levels (25.94% of control vs 27.15% of UV10) and the inhibition of shade avoidance syndrome (denser growth with shorter stolons and internodes) were only observed in offspring of high UV-B parents. This difference was related to the eustress and stress effects of low and high UV-B, respectively. Overall, the behaviour of D. indica under heterogeneous light conditions was regulated by the parental UV-B exposure. Moreover, certain performance improvements helped offspring pre-regulate growth to cope with future environments and were probably associated with the effects of maternal DNA methylation variations in UV-B-radiated parents.

Allometric relationships, functional differentiations, and scaling of growth rates across 151 tree species in China

AbstractChinese trees adapted to wet, warm, highly heterogeneous, and dynamic light environments might be expected to systematically differ from other non‐Chinese trees based on stem allometry and life em allo. However, our understanding of the extent to which how differentiation in forest architecture and species size influences tree allometry relates to fundamental physiological and ecological trade‐offs, climate, forest structure, and function is limited. We quantified height–diameter allometries and growth increments of 13,594 individuals of 151 tree species across China. Although trees grew statistically indistinguishably from the theoretically predicted exponent, the height–diameter allometries for trees in China deviated from tree patterns in other continental forests worldwide. Chinese trees showed substantially lower heights at a given diameter for larger trees and asymptotic heights (33.05 m), with values were 5.5% to 43.3% lower than those estimated for continental rain forest trees in South America, Australia, Southeast Asia, and Africa. In total, 149 of 151 species showed significant nonlinearity of height–diameter allometry with asymptotic heights ranging from 6.33 to 50.54 m; additionally, half (74/149 species) fell in the range of 20–33.5 m, averaging 21.98 ± 8.70 m (SE) for canopy trees. In subsets of canopy trees sampled from larger‐statured species and deciduous trees, the approximated maximum heights were 14–42% larger than those of smaller‐statured and evergreens and those in other wet and warmer regions found worldwide. These results suggest flexible phenotypic responses of height–diameter allometries to heterogeneous light conditions along understory‐canopy gradients. Average growth rates, varying from 0.02 to 0.78 cm/yr (mean of 0.25 cm/yr), were positively related to the asymptotic height, diameter at inflection, and allometric intercept but negatively correlated with the shape of the asymptotic curve and allometric slope. Species with asymptotic heights above 30 m or inflection sizes larger than 20 cm clearly depicted higher growth rates than smaller ones.

Effects of clonal integration on allelopathy of invasive plantWedelia trilobataunder heterogeneous light conditions

AbstractPlant invasion is one of the most serious threats to ecosystems worldwide. When invasive plants with the ability of clonal growth invading or colonizing in new habitat, their interconnected ramets may suffer from heterogeneous light. Effects of clonal integration on allelopathy of invasive plants are poorly understood under heterogeneous light conditions. To investigate the effects of clonal integration on allelopathy of invasive plant Wedelia trilobata under heterogeneous light conditions, a pot experiment was conducted by using its clonal fragments with two successive ramets. The older ramets were exposed to full light, whereas the younger ones were subjected to 20% full light. The younger ramets of each clonal fragment were adjacently grown with a target plant (one tomato seedling) in a pot. Stolon between two successive ramets was either severed or retained intact. In addition, two tomato seedlings (one as target plant) were adjacently grown in a pot as contrast. Compared with severing stolon, biomass accumulation, foliar chlorophyll and nitrogen contents, chlorophyll fluorescence parameters and net photosynthetic rates of the target plants as well as their root length and activity, were significantly decreased when stolon between interconnected ramets of W. trilobata retained intact. Under heterogeneous light conditions, transportation or sharing of carbohydrate between two successive ramets enhanced allelopathy of the young ramets subjected to 20% full light treatment. It is suggested that clonal integration may be important for invasion or colonization of invasive plants with ability of clonal growth under heterogeneous light conditions.

Heterogeneous Light Conditions Reduce the Assimilate Translocation Towards Maize Ears.

The border row crop in strip intercropped maize is often exposed to heterogeneous light conditions, resulting in increased photosynthesis and yield decreased. Previous studies have focused on photosynthetic productivity, whereas carbon allocation could also be one of the major causes of decreased yield. However, carbon distribution remains unclear in partially shaded conditions. In the present study, we applied heterogeneous light conditions (T), and one side of plants was shaded (T-30%), keeping the other side fully exposed to light (T-100%), as compared to control plants that were exposed entirely to full-light (CK). Dry weight, carbon assimilation, 13C abundance, and transport tissue structure were analyzed to clarify the carbon distribution in partial shading of plants. T caused a marked decline in dry weight and harvest index (HI), whereas dry weight in unshaded and shaded leaves did not differ. Net photosynthesis rate (Pn), the activity of sucrose phosphate synthase enzymes (SPS), and sucrose concentration increased in unshaded leaves. Appropriately, 5.7% of the 13C from unshaded leaves was transferred to shaded leaves. Furthermore, plasmodesma density in the unshaded (T-100%) and shaded (T-30%) leaves in T was not significantly different but was lower than that of CK. Similarly, the vascular bundle total area of T was decreased. 13C transfer from unshaded leaves to ear in T was decreased by 18.0% compared with that in CK. Moreover, 13C and sucrose concentration of stem in T were higher than those in CK. Our results suggested that, under heterogeneous light, shaded leaves as a sink imported the carbohydrates from the unshaded leaves. Ear and shaded leaf competed for carbohydrates, and were not conducive to tissue structure of sucrose transport, resulting in a decrease in the carbon proportion in the ear, harvest index, and ear weight.

Plasticity and light sensitivity of leaf hydraulic conductance to fast changes in irradiance in common hazel (Corylus avellana L.)

Forest understory species have to acclimatize to highly heterogeneous light conditions inside forest canopies in order to utilize available resources efficiently. Light sensitivity and response speed of hydraulic conductance (KL) of common hazel (Corylus avellana L.) to fast changes in irradiance was studied in leaves from three different growth light conditions–sun-exposed, moderate shade, and deep shade. The KL of sun-exposed leaves was approximately 3-fold higher when compared to deep-shade leaves, indicating a strong dependence of leaf hydraulic capacity on light conditions. The KL of sun-exposed leaves increased by a factor of nearly four from minimal values recorded in darkness to maximal values in high light compared to deep-shade leaves. Reaction speed of KL to reach maximum values in response to light was nearly five times higher for sun-exposed vs deep-shade leaves. Plasticity indices of KL for sun-exposed and deep-shade leaves were 0.44 and 0.27, respectively. Higher light sensitivity enables a faster and more plastic response of KL to variable light conditions in sun leaves and enhances the ability of plants to maximize resource utilization under more beneficial environmental conditions.

Estimating below-canopy light regimes using airborne laser scanning: An application to plant community analysis.

Light is a key driver of forest biodiversity and functioning. Light regimes beneath tree canopies are mainly driven by the solar angle, topography, and vegetation structure, whose three‐dimensional complexity creates heterogeneous light conditions that are challenging to quantify, especially across large areas. Remotely sensed canopy structure data from airborne laser scanning (ALS) provide outstanding opportunities for advancement in this respect. We used ALS point clouds and a digital terrain model to produce hemispherical photographs from which we derived indices of nondirectional diffuse skylight and direct sunlight reaching the understory. We validated our approach by comparing the performance of these indices, as well as canopy closure (CCl) and canopy cover (CCo), for explaining the light conditions experienced by forest plant communities, as indicated by the Landolt indicator values for light (L light) from 43 vegetation surveys along an elevational gradient. We applied variation partitioning to analyze how the independent and joint statistical effects of light, macroclimate, and soil on the spatial variation in plant species composition (i.e., turnover, Simpson dissimilarity, β SIM) depend on light approximation methodology. Diffuse light explained L light best, followed by direct light, CCl and CCo (R2 = .31, .23, .22, and .22, respectively). The combination of diffuse and direct light improved the model performance for β SIM compared with CCl and CCo (R2 = .30, .27 and .24, respectively). The independent effect of macroclimate on β SIM dropped from an R 2 of .15 to .10 when diffuse light and direct light were included. The ALS methods presented here outperform conventional approximations of below‐canopy light conditions, which can now efficiently be quantified along entire horizontal and vertical forest gradients, even in topographically complex environments such as mountains. The effect of macroclimate on forest plant communities is prone to be overestimated if local light regimes and associated microclimates are not accurately accounted for.

Involvement of carbohydrates in long-term light-dependent systemic regulation on photosynthesis of maize under light heterogeneity

ABSTRACTIt is widely recognized that different parts of a plant can communicate with each other via light-dependent long-distance signaling under heterogeneous light conditions. However, the mechanism of such systemic signaling has not been revealed yet. Our studies on different species suggest the involvement of carbohydrates in light-dependent systemic regulation between different parts of a plant under both short- and long-term light heterogeneity. Leaves exposed to better light condition perform enhanced photosynthetic capacity, and act to compensate for the decline in photosynthesis of other leaves under bad light condition within the same plant. This kind of compensatory photosynthesis has a close relationship to the distribution of carbohydrates, and can be regarded as an integrative strategy to make efficient use of sunlight at the whole-plant level.

Light-induced systemic signalling down-regulates photosynthetic performance of soybean leaves with different directional effects.

When plants are exposed to a heterogeneous environment, photosynthesis of leaves is not only determined by their local condition, but also by certain signals from other parts of the same plant, termed systemic regulation. Our present study was conducted to investigate the effects of light-dependent systemic regulation on the photosynthetic performance of soybean (Glycine max L. Merr.) under heterogeneous light conditions. Soybean plants were treated with heterogeneous light. Then gas exchange characteristics were measured to evaluate the photosynthetic performance of leaves. Parameters related to photosynthetic pigments, chlorophyll fluorescence, Rubisco and photosynthates were examined to study the mechanisms of light-dependent systemic regulation on photosynthesis. Light-induced systemic signalling by illuminated leaves reduced the Pn of both upper and lower non-illuminated leaves on the same soybean plant. The decrease in gs and increase in Ci in these non-illuminated leaves indicated restriction of carbon assimilation, which was further verified by the decline in content and activity of Rubisco. However, the activation state of Rubisco decreased only in upper non-illuminated leaves. Quantum efficiency of PSII (ΦPSII) and ETR also decreased only in upper non-illuminated leaves. Moreover, the effects of light-induced systemic signalling on carbohydrate content were also detectable only in upper non-illuminated leaves. Light-induced systemic signalling by illuminated leaves restricts carbon assimilation and down-regulates photosynthetic performance of non-illuminated leaves within a soybean plant. However, effects of such systemic regulation differed when regulated in upward or downward direction.

<b>Leaf architecture and symmetry of understory tree species of an Araucaria forest

Shade plants of ombrophilous forests are subjected to light-limiting conditions and need to invest in architectural structures associated with leaf symmetry to increase light capture. This study investigated the leaf architecture of six Araucaria forest tree species with distinct symmetry: Cupania vernalis, Casearia sylvestris, Schinus terebinthifolius, Piper gaudichaudianum, Roupala brasiliensis and Cedrela fissilis . We hypothesized that symmetry, associated with other traits, minimizes self-shading. Asymmetry index, petiole length, total leaf area, leaf angle, internode length and stem diameter were measured. The asymmetry index did not indicate a clear distinction between asymmetric and symmetric leaves. Leaves classified as asymmetric had higher values for the asymmetry index in the median and basal regions of the leaf, while symmetrical leaves had higher values in the apical region. The results also indicated an adjustment among structural leaf traits that facilitated a three-dimensional organization that produced an advantageous arrangement for light capture, which seems to be a response to selective pressure by the heterogeneous light conditions of the ombrophilous forest understory.

Variation in species light acquisition traits under fluctuating light regimes: implications for non‐equilibrium coexistence

Resource distribution heterogeneity offers niche opportunities for species with different functional traits to develop and potentially coexist. Available light (photosynthetically active radiation or PAR) for suspended algae (phytoplankton) may fluctuate greatly over time and space. Species‐specific light acquisition traits capture important aspects of the ecophysiology of phytoplankton and characterize species growth at either limiting or saturating daily PAR supply. Efforts have been made to explain phytoplankton coexistence using species‐specific light acquisition traits under constant light conditions, but not under fluctuating light regimes that should facilitate non‐equilibrium coexistence. In the well‐mixed, hypertrophic Lake TaiHu (China), we incubated the phytoplankton community in bottles placed either at fixed depths or moved vertically through the water column to mimic vertical mixing. Incubations at constant depths received only the diurnal changes in light, while the moving bottles received rapidly fluctuating light. Species‐specific light acquisition traits of dominant cyanobacteria (Anabaena flos‐aquae, Microcystis spp.) and diatom (Aulacoseira granulata, Cyclotella pseudostelligera) species were characterized from their growth–light relationships that could explain relative biomasses along the daily PAR gradient under both constant and fluctuating light. Our study demonstrates the importance of interspecific differences in affinities to limiting and saturating light for the coexistence of phytoplankton species in spatially heterogeneous light conditions. Furthermore, we observed strong intraspecific differences in light acquisition traits between incubation under constant and fluctuating light – leading to the reversal of light utilization strategies of species. This increased the niche space for acclimated species, precluding competitive exclusion. These observations could enhance our understanding of the mechanisms behind the Paradox of the Plankton.

A Robust Face Recognition Method Combining LBP with Multi-mirror Symmetry for Images with Various Face Interferences

Face recognition (FR) is a practical application of pattern recognition (PR) and remains a compelling topic in the study of computer vision. However, in real-world FR systems, interferences in images, including illumination condition, occlusion, facial expression and pose variation, make the recognition task challenging. This study explored the impact of those interferences on FR performance and attempted to alleviate it by taking face symmetry into account. A novel and robust FR method was proposed by combining multi-mirror symmetry with local binary pattern (LBP), namely multi-mirror local binary pattern (MMLBP). To enhance FR performance with various interferences, the MMLBP can 1) adaptively compensate lighting under heterogeneous lighting conditions, and 2) generate extracted image features that are much closer to those under well-controlled conditions (i.e., frontal facial images without expression). Therefore, in contrast with the later variations of LBP, the symmetrical singular value decomposition representation (SSVDR) algorithm utilizing the facial symmetry and a state-of-art non-LBP method, the MMLBP method is shown to successfully handle various image interferences that are common in FR applications without preprocessing operation and a large number of training images. The proposed method was validated with four public data sets. According to our analysis, the MMLBP method was demonstrated to achieve robust performance regardless of image interferences.

Effect of clonal integration on nitrogen cycling in rhizosphere of rhizomatous clonal plant, Phyllostachys bissetii, under heterogeneous light

Clonal integration plays an important role in clonal plant adapting to heterogeneous habitats. It was postulated that clonal integration could exhibit positive effects on nitrogen cycling in the rhizosphere of clonal plant subjected to heterogeneous light conditions. An in-situ experiment was conducted using clonal fragments of Phyllostachys bissetii with two successive ramets. Shading treatments were applied to offspring or mother ramets, respectively, whereas counterparts were treated to full sunlight. Rhizomes between two successive ramets were either severed or connected. Extracellular enzyme activities and nitrogen turnover were measured, as well as soil properties. Abundance of functional genes (archaeal or bacterial amoA, nifH) in the rhizosphere of shaded, offspring or mother ramets were determined using quantitative polymerase chain reaction. Carbon or nitrogen availabilities were significantly influenced by clonal integration in the rhizosphere of shaded ramets. Clonal integration significantly increased extracellular enzyme activities and abundance of functional genes in the rhizosphere of shaded ramets. When rhizomes were connected, higher nitrogen turnover (nitrogen mineralization or nitrification rates) was exhibited in the rhizosphere of shaded offspring ramets. However, nitrogen turnover was significantly decreased by clonal integration in the rhizosphere of shaded mother ramets. Path analysis indicated that nitrogen turnover in the rhizosphere of shaded, offspring or mother ramets were primarily driven by the response of soil microorganisms to dissolved organic carbon or nitrogen. This unique in-situ experiment provided insights into the mechanism of nutrient recycling mediated by clonal integration. It was suggested that effects of clonal integration on the rhizosphere microbial processes were dependent on direction of photosynthates transport in clonal plant subjected to heterogeneous light conditions.

Invasive alien plants benefit more from clonal integration in heterogeneous environments than natives.

What confers invasive alien plants a competitive advantage over native plants remains open to debate. Many of the world's worst invasive alien plants are clonal and able to share resources within clones (clonal integration), particularly in heterogeneous environments. Here, we tested the hypothesis that clonal integration benefits invasive clonal plants more than natives and thus confers invasives a competitive advantage. We selected five congeneric and naturally co-occurring pairs of invasive alien and native clonal plants in China, and grew pairs of connected and disconnected ramets under heterogeneous light, soil nutrient and water conditions that are commonly encountered by alien plants during their invasion into new areas. Clonal integration increased biomass of all plants in all three heterogeneous resource environments. However, invasive plants benefited more from clonal integration than natives. Consequently, invasive plants produced more biomass than natives. Our results indicate that clonal integration may confer invasive alien clonal plants a competitive advantage over natives. Therefore, differences in the ability of clonal integration could potentially explain, at least partly, the invasion success of alien clonal plants in areas where resources are heterogeneously distributed.

Nitrogen retention and loss in unfertilized lawns across a light gradient

A desirable function for terrestrial urban ecosystems is the mitigation of nitrogen (N) pollution associated with cities and suburbs. To assist in maximizing this function, identifying locations of sources and sinks of N in the urban environment is crucial to inform management strategies. Lawns are an extensive land cover in urbanized landscapes, and in general, they have demonstrated the capacity to function as a sink for N inputs. How N is cycled by lawns, however, is likely not uniform across the physical heterogeneity or management activities that exist in lawns. We investigated the influence of heterogeneity in light availability on N cycling in lawns that were irrigated but not fertilized. Light availability is affected by tree canopy and built structures and is, therefore, heterogeneous both within individual lawns and among lawns. Light is expected to control N retention and loss through effects on primary productivity. We experimentally examined N regulation over one calendar year by measuring net primary production (NPP), N retention using an isotopic tracer, and N leaching in existing unfertilized lawns under heterogeneous light conditions. We used a budgetary approach to estimate gaseous N loss which we assume is primarily via denitrification. Light functioned as a limiting resource for primary productivity. From low to high light conditions, annual NPP increased 177 to 430 g C∙m−2∙y−1 and retention of N isotope tracer increased from 50 to 65% as a result of increased retention in plants. Nitrate leaching losses were low overall and were not affected by light levels. Light availability regulated the fate of N inputs and unfertilized lawns may function as substantial sinks for reactive N through storage in the terrestrial system and N loss by denitrification. However, whether or not denitrification is generally an N sink will depend on the ratio of non-reactive (i.e., N2) to reactive (i.e., N2O, NO) denitrification products. Overall, we find that effective strategies for managing N sources and sinks in cities will likely need to consider light availability, particularly in systems receiving water subsidies via irrigation.

Leaf diseases drive the Janzen-Connell mechanism regardless of light conditions: a 3-year field study.

In forests, negative density/distance-dependent seedling mortality (NDD) caused by natural enemies plays a key role in maintaining species diversity [Janzen-Connell (J-C) model]. However, the relative importance of natural enemies in mediating NDD under heterogeneous light conditions has remained unclear. We examined the relative importance of pathogens (i.e., soil pathogens, leaf diseases) on seedling performance in forest understories (FUs) and gaps (gaps) during a 3-year period (results of first year of our study have been previously reported). For the hardwood, Prunus grayana, we investigated seedling mortality, morbidity agents, growth, and root infection by arbuscular mycorrhizal fungi (AMF) beneath conspecific and heterospecific adults in FUs and gaps. Seedling mortality was higher beneath conspecific than heterospecific adults throughout 3years at both sites, mainly due to continuous leaf disease (i.e., angular leaf spot), whereas damping-off diseases caused mortality only in the first year. Beneath each adult, seedling mortality was higher in FUs than in gaps until second year, but it did not differ between two habitat types in the third year, because leaf diseases caused severe damage even in gaps. Seedling mass was significantly lower beneath conspecific adults. AMF infection of seedlings was also lower beneath conspecific adults, while it was higher in gaps than in FUs beneath both adults. This study demonstrates that the J-C model in a hardwood tree, P. grayana is mainly driven by high NDD seedling mortality caused by airborne leaf diseases, which continuously attack seedlings in a NDD manner regardless of environmental light conditions.

Congruence of the spatial pattern of light and understory vegetation in an old-growth, temperate mixed forest

Light is one of the most important drivers of understory vegetation in forests, influencing the patterns of total cover as well as the abundance of individual species.Based on a multi-scale approach, the relationships between the amount and pattern of relative diffuse light and forest understory were studied in an old-growth, temperate mixed forest (Hungary). The recorded vegetation variables were the cover of the vascular understory (herbs, woody seedlings), the bryophyte layer, and some selected vascular understory species.The pattern of light showed aggregations at two scales: 10×10 and 25×25m. Both vascular understory and bryophyte cover had significant positive correlations with light availability, and their spatial pattern was related to it. The pattern of seedlings displayed the strongest relationships with that of light at a coarser scale (25×25m) than herbs and bryophytes (10×10m). At the species level, Festuca heterophylla, Fragaria vesca and Poa nemoralis were characterized as light-demanding herbaceous species (their spatial pattern was congruent with light), Brachypodium sylvaticum and Carex pallescens were transitional, while some species proved to be shade-tolerant (e.g. Ajuga reptans, Dryopteris carthusiana, Viola reichenbachiana). Regarding seedlings, the patterns of Betula pendula, Carpinus betulus, Pinus sylvestris and Quercus petraea were related to the pattern of light.According to our observations, diversity and composition of vascular forest understory and bryophytes were related to heterogeneous light conditions. Forest management should maintain continuous shelter on the stand level; however, smaller gaps are necessary for the survival of light-demanding forest herbs and bryophytes, and larger gaps for tree seedlings.

Environmental drivers of the composition and diversity of the herb layer in mixed temperate forests in Hungary

Herbaceous understory vegetation is an important part of temperate forested ecosystems, the diversity and composition of which are strongly dependent on the conditions of the forest stand and the landscape. The aim of this study was to find the most important environmental drivers influencing understory herb layer species composition (explored with multivariate analysis), and richness and cover (analysed by linear modelling) in managed mixed forests in West Hungary. Our detailed inventory showed that the most important factors increasing the diversity and cover of the understory are light, tree species richness, and landscape diversity. Composition is also mainly influenced by light conditions and tree species richness, with minor effects of tree species composition, soil texture, and moss cover. As the strongest influencing factors are closely linked to stand structure and tree species composition, they can either directly or indirectly be altered by forest management. In the studied region, heterogeneous light conditions and canopy structure, the maintenance of tree species richness and forest continuity are key elements for the conservation of forest herbs. Forestry that maintains continuous forest cover and the tree selection management system can better provide these conditions than the presently widely used shelterwood management system.