Biomass partitioning and growth efficiency in four naturally regenerated forest tree species

Current forest growth models and yield tables are almost exclusively based on data from mature trees, reducing their applicability to young and developing stands. To address this gap, young European beech, sessile oak, Scots pine and Norway spruce trees approximately 0 to 10 years old were destructively sampled in a range of naturally regenerated forest stands in Central Europe. Diameter at base and height were first measured in situ for up to 175 individuals per species. Subsequently, the trees were excavated and dry biomass of foliage, branches, stems and roots was measured. Allometric relations were then used to calculate biomass allocation coefficients (BAC) and growth efficiency (GE) patterns in young trees. We found large differences in BAC and GE between broadleaves and conifers, but also between species within these categories. Both BAC and GE are strongly age-specific in young trees, their rapidly changing values reflecting different growth strategies in the earliest stages of growth. We show that linear relationships describing biomass allocation in older trees are not applicable in young trees. To accurately predict forest biomass and carbon stocks, forest growth models need to include species and age specific parameters of biomass allocation patterns.

A novel method for assessing risks to pollinators from plant protection products using honeybees as a model species

Pollination is one of the most important ecosystem services in agroecosystems and supports food production. Pollinators are potentially at risk being exposed to pesticides and the main route of exposure is direct contact, in some cases ingestion, of contaminated materials such as pollen, nectar, flowers and foliage. To date there are no suitable methods for predicting pesticide exposure for pollinators, therefore official procedures to assess pesticide risk are based on a Hazard Quotient. Here we develop a procedure to assess exposure and risk for pollinators based on the foraging behaviour of honeybees (Apis mellifera) and using this species as indicator representative of pollinating insects. The method was applied in 13 European field sites with different climatic, landscape and land use characteristics. The level of risk during the crop growing season was evaluated as a function of the active ingredients used and application regime. Risk levels were primarily determined by the agronomic practices employed (i.e. crop type, pest control method, pesticide use), and there was a clear temporal partitioning of risks through time. Generally the risk was higher in sites cultivated with permanent crops, such as vineyard and olive, than in annual crops, such as cereals and oil seed rape. The greatest level of risk is generally found at the beginning of the growing season for annual crops and later in June–July for permanent crops.

Genetic diversity of red-fleshed apples (Malus)

Anthocyanins are flavonoid pigments imparting red, blue, or purple pigmentation to fruits, flowers and foliage. These compounds are powerful antioxidants in vitro, and are widely believed to contribute to human health. The fruit of the domestic apple (Malus x domestica) is a popular and important source of nutrients, and is considered one of the top ‘functional foods’—those foods that have inherent health-promoting benefits beyond basic nutritional value. The pigmentation of typical red apple fruits results from accumulation of anthocyanin in the skin. However, numerous genotypes of Malus are known that synthesize anthocyanin in additional fruit tissues including the core and cortex (flesh). Red-fleshed apple genotypes are an attractive starting point for development of novel varieties for consumption and nutraceutical use through traditional breeding and biotechnology. However, cultivar development is limited by lack of characterization of the diversity of genetic backgrounds showing this trait. We identified and cataloged red-fleshed apple genotypes from four Malus diversity collections representing over 3,000 accessions including domestic cultivars, wild species, and named hybrids. We found a striking range of flesh color intensity and pattern among accessions, including those carrying the MYB10 R 6 allele conferring ectopic expression of a key transcriptional regulator of anthocyanin biosynthesis. Although MYB10 R 6 was strongly associated with red-fleshed fruit among genotypes, this allele was neither sufficient nor required for this trait in all genotypes. Nearly all red-fleshed accessions tested could be traced back to ‘Niedzwetzkyana’, a presumed natural form of M. sieversii native to central Asia.

The REFLEX project: Comparing different algorithms and implementations for the inversion of a terrestrial ecosystem model against eddy covariance data

We describe a model-data fusion (MDF) inter-comparison project (REFLEX), which compared various algorithms for estimating carbon (C) model parameters consistent with both measured carbon fluxes and states and a simple C model. Participants were provided with the model and with both synthetic net ecosystem exchange (NEE) of CO2 and leaf area index (LAI) data, generated from the model with added noise, and observed NEE and LAI data from two eddy covariance sites. Participants endeavoured to estimate model parameters and states consistent with the model for all cases over the two years for which data were provided, and generate predictions for one additional year without observations. Nine participants contributed results using Metropolis algorithms, Kalman filters and a genetic algorithm. For the synthetic data case, parameter estimates compared well with the true values. The results of the analyses indicated that parameters linked directly to gross primary production (GPP) and ecosystem respiration, such as those related to foliage allocation and turnover, or temperature sensitivity of heterotrophic respiration, were best constrained and characterised. Poorly estimated parameters were those related to the allocation to and turnover of fine root/wood pools. Estimates of confidence intervals varied among algorithms, but several algorithms successfully located the true values of annual fluxes from synthetic experiments within relatively narrow 90% confidence intervals, achieving >80% success rate and mean NEE confidence intervals <110 gC m−2 year−1 for the synthetic case. Annual C flux estimates generated by participants generally agreed with gap-filling approaches using half-hourly data. The estimation of ecosystem respiration and GPP through MDF agreed well with outputs from partitioning studies using half-hourly data. Confidence limits on annual NEE increased by an average of 88% in the prediction year compared to the previous year, when data were available. Confidence intervals on annual NEE increased by 30% when observed data were used instead of synthetic data, reflecting and quantifying the addition of model error. Finally, our analyses indicated that incorporating additional constraints, using data on C pools (wood, soil and fine roots) would help to reduce uncertainties for model parameters poorly served by eddy covariance data.

The effects of a changing pollution climate on throughfall deposition and cycling in a forested area in southern England

This study compares two sets of measurements of the composition of bulk precipitation and throughfall at a site in southern England with a 20-year gap between them. During this time, SO2 emissions from the UK fell by 82%, NOx emissions by 35% and NH3 emissions by 7%. These reductions were partly reflected in bulk precipitation, with deposition reductions of 56% in SO4,38% in NO3, 32% in NH4, and 73% in H+. In throughfall under Scots pine, the effects were more dramatic, with an 89% reduction in SO4 deposition and a 98% reduction in H+ deposition. The mean pH under these trees increased from 2.85 to 4.30. Nitrate and ammonium deposition in throughfall increased slightly, however. In the earlier period, the Scots pines were unable to neutralise the high flux of acidity associated with sulphur deposition, even though this was not a highly polluted part of the UK, and deciduous trees (oak and birch) were only able to neutralise it in summer when the leaves were present. In the later period, the sulphur flux had reduced to the point where the acidity could be neutralised by all species — the neutralisation mechanism is thus likely to be largely leaching of base cations and buffering substances from the foliage. The high fluxes are partly due to the fact that these are 60–80 year old trees growing in an open forest structure. The increase in NO3 and NH4 in throughfall in spite of decreased deposition seems likely due to a decrease in foliar uptake, perhaps due to the increasing nitrogen saturation of the catchment soils. These changes may increase the rate of soil microbial activity as nitrogen increases and acidity declines, with consequent effects on water quality of the catchment drainage stream.

Climate change impacts on crop productivity in Africa and South Asia

Climate change is a serious threat to crop productivity in regions that are already food insecure. We assessed the projected impacts of climate change on the yield of eight major crops in Africa and South Asia using a systematic review and meta-analysis of data in 52 original publications from an initial screen of 1144 studies. Here we show that the projected mean change in yield of all crops is − 8% by the 2050s in both regions. Across Africa, mean yield changes of − 17% (wheat), − 5% (maize), − 15% (sorghum) and − 10% (millet) and across South Asia of − 16% (maize) and − 11% (sorghum) were estimated. No mean change in yield was detected for rice. The limited number of studies identified for cassava, sugarcane and yams precluded any opportunity to conduct a meta-analysis for these crops. Variation about the projected mean yield change for all crops was smaller in studies that used an ensemble of > 3 climate (GCM) models. Conversely, complex simulation studies that used biophysical crop models showed the greatest variation in mean yield changes. Evidence of crop yield impact in Africa and South Asia is robust for wheat, maize, sorghum and millet, and either inconclusive, absent or contradictory for rice, cassava and sugarcane.

Acetone enhances the direct analysis of procyanidin- and prodelphinidin-based condensed tannins in lotus species by the butanol-HCl-iron assay

The butanol-HCl spectrophotometric assay is widely used for quantifying extractable and insoluble condensed tannins (CT, syn. proanthocyanidins) in foods, feeds, and foliage of herbaceous and woody plants, but the method underestimates total CT content when applied directly to plant material. To improve CT quantitation, we tested various cosolvents with butanol-HCl and found that acetone increased anthocyanidin yields from two forage Lotus species having contrasting procyanidin and prodelphinidin compositions. A butanol-HCl-iron assay run with 50% (v/v) acetone gave linear responses with Lotus CT standards and increased estimates of total CT in Lotus herbage and leaves by up to 3.2-fold over the conventional method run without acetone. The use of thiolysis to determine the purity of CT standards further improved quantitation. Gel-state 13C and 1H–13C HSQC NMR spectra of insoluble residues collected after butanol-HCl assays revealed that acetone increased anthocyanidin yields by facilitating complete solubilization of CT from tissue.

Replacing Norway spruce with European beech: a comparison of biomass and Net Primary Production patterns in young stands

An alteration of species composition in temperate forests – both managed and natural - is one of the expected effects of environmental change. Present forest tree species ranges will be altered by changing environmental conditions. By a combination of continuous and destructive sampling, we compared biomass stocks and annual NPP in naturally regenerated stands of Norway spruce and European beech. We purposely selected a site where future environmental conditions are predicted to favour beech over presently dominant spruce. We found no difference in overall productivity, but biomass allocation differed significantly between the two species. Beech allocated more assimilates to stem and roots than spruce. There was no significant difference between the species in NPP of the fast turnover biomass pool comprising foliage and fine roots. Maximum height growth occurred about a month earlier than in spruce, potentially changing the timing of carbon (C) flow into the soil pools. We show that the replacement of spruce by beech will result in changes in forest biomass allocation and in alterations of belowground C cycle. Such changes will affect forest ecosystem function by modifying the magnitude and timing of certain C fluxes, but also by potentially changing the species composition of forest biota dependent on them.

Simulation of tree-ring widths with a model for primary production, carbon allocation, and growth

We present a simple, generic model of annual tree growth, called "T". This model accepts input from a first-principles light-use efficiency model (the "P" model). The P model provides values for gross primary production (GPP) per unit of absorbed photosynthetically active radiation (PAR). Absorbed PAR is estimated from the current leaf area. GPP is allocated to foliage, transport tissue, and fine-root production and respiration in such a way as to satisfy well-understood dimensional and functional relationships. Our approach thereby integrates two modelling approaches separately developed in the global carbon-cycle and forest-science literature. The T model can represent both ontogenetic effects (the impact of ageing) and the effects of environmental variations and trends (climate and CO2) on growth. Driven by local climate records, the model was applied to simulate ring widths during the period 1958–2006 for multiple trees of Pinus koraiensis from the Changbai Mountains in northeastern China. Each tree was initialised at its actual diameter at the time when local climate records started. The model produces realistic simulations of the interannual variability in ring width for different age cohorts (young, mature, and old). Both the simulations and observations show a significant positive response of tree-ring width to growing-season total photosynthetically active radiation (PAR0) and the ratio of actual to potential evapotranspiration (α), and a significant negative response to mean annual temperature (MAT). The slopes of the simulated and observed relationships with PAR0 and α are similar; the negative response to MAT is underestimated by the model. Comparison of simulations with fixed and changing atmospheric CO2 concentration shows that CO2 fertilisation over the past 50 years is too small to be distinguished in the ring-width data, given ontogenetic trends and interannual variability in climate.

Validation of Canopy Height Profile methodology for small-footprint full-waveform airborne LiDAR data in a discontinuous canopy environment

A Canopy Height Profile (CHP) procedure presented in Harding et al. (2001) for large footprint LiDAR data was tested in a closed canopy environment as a way of extracting vertical foliage profiles from LiDAR raw-waveform. In this study, an adaptation of this method to small-footprint data has been shown, tested and validated in an Australian sparse canopy forest at plot- and site-level. Further, the methodology itself has been enhanced by implementing a dataset-adjusted reflectance ratio calculation according to Armston et al. (2013) in the processing chain, and tested against a fixed ratio of 0.5 estimated for the laser wavelength of 1550nm. As a by-product of the methodology, effective leaf area index (LAIe) estimates were derived and compared to hemispherical photography-derived values. To assess the influence of LiDAR aggregation area size on the estimates in a sparse canopy environment, LiDAR CHPs and LAIes were generated by aggregating waveforms to plot- and site-level footprints (plot/site-aggregated) as well as in 5m grids (grid-processed). LiDAR profiles were then compared to leaf biomass field profiles generated based on field tree measurements. The correlation between field and LiDAR profiles was very high, with a mean R2 of 0.75 at plot-level and 0.86 at site-level for 55 plots and the corresponding 11 sites. Gridding had almost no impact on the correlation between LiDAR and field profiles (only marginally improvement), nor did the dataset-adjusted reflectance ratio. However, gridding and the dataset-adjusted reflectance ratio were found to improve the correlation between raw-waveform LiDAR and hemispherical photography LAIe estimates, yielding the highest correlations of 0.61 at plot-level and of 0.83 at site-level. This proved the validity of the approach and superiority of dataset-adjusted reflectance ratio of Armston et al. (2013) over a fixed ratio of 0.5 for LAIe estimation, as well as showed the adequacy of small-footprint LiDAR data for LAIe estimation in discontinuous canopy forests.

A model analysis of climate and CO2 controls on tree growth in a semi-arid woodland

We used a light-use efficiency model of photosynthesis coupled with a dynamic carbon allocation and tree-growth model to simulate annual growth of the gymnosperm Callitris columellaris in the semi-arid Great Western Woodlands, Western Australia, over the past 100 years. Parameter values were derived from independent observations except for sapwood specific respiration rate, fine-root turnover time, fine-root specific respiration rate and the ratio of fine-root mass to foliage area, which were estimated by Bayesian optimization. The model reproduced the general pattern of interannual variability in radial growth (tree-ring width), including the response to the shift in precipitation regimes that occurred in the 1960s. Simulated and observed responses to climate were consistent. Both showed a significant positive response of tree-ring width to total photosynthetically active radiation received and to the ratio of modeled actual to equilibrium evapotranspiration, and a significant negative response to vapour pressure deficit. However, the simulations showed an enhancement of radial growth in response to increasing atmospheric CO2 concentration (ppm) ([CO2]) during recent decades that is not present in the observations. The discrepancy disappeared when the model was recalibrated on successive 30-year windows. Then the ratio of fine-root mass to foliage area increases by 14% (from 0.127 to 0.144 kg C m-2) as [CO2] increased while the other three estimated parameters remained constant. The absence of a signal of increasing [CO2] has been noted in many tree-ring records, despite the enhancement of photosynthetic rates and water-use efficiency resulting from increasing [CO2]. Our simulations suggest that this behaviour could be explained as a consequence of a shift towards below-ground carbon allocation.

Ectomycorrhizal symbiosis can enhance plant nutrition through improved access to discrete organic nutrient patches of high resource quality

It is known that roots can respond to patches of fertility; however, root proliferation is often too slow to exploit resources fully, and organic nutrient patches may be broken down and leached, immobilized or chemically fixed before they are invaded by the root system. The ability of fungal hyphae to exploit resource patches is far greater than that of roots due to their innate physiological and morphological plasticity, which allows comprehensive exploration and rapid colonization of resource patches in soils. The fungal symbionts of ectomycorrhizal plants excrete significant quantities of enzymes such as chitinases, phosphatases and proteases. These might allow the organic residue to be tapped directly for nutrients such as N and P. Pot experiments conducted with nutrient-stressed ectomycorrhizal and control willow plants showed that when high quality organic nutrient patches were added, they were colonized rapidly by the ectomycorrhizal mycelium. These established willows (0.5 m tall) were colonized by Hebeloma syrjense P. Karst. for 1 year prior to nutrient patch addition. Within days after patch addition, colour changes in the leaves of the mycorrhizal plants (reflecting improved nutrition) were apparent, and after I month the concentration of N and P in the foliage of mycorrhizal plants was significantly greater than that in non-mycorrhizal plants subject to the same nutrient addition. It seems likely that the mycorrhizal plants were able to compete effectively with the wider soil microbiota and tap directly into the high quality organic resource patch via their extra-radical mycelium. We hypothesize that ectomycorrhizal plants may reclaim some of the N and P invested in seed production by direct recycling from failed seeds in the soil. The rapid exploitation of similar discrete, transient, high-quality nutrient patches may have led to underestimations when determining the nutritional benefits of ectomycorrhizal colonization.

Specific leaf area and leaf area index in developing stands of Fagus sylvatica L. and Picea abies Karst

European beech (Fagus sylvatica L.) and Norway spruce (Picea abies Karst.) are two of the most ecologically and economically important forest tree species in Europe. These two species co-occur in many locations in Europe, leading to direct competition for canopy space. Foliage characteristics of two naturally regenerated pure stands of beech and spruce with fully closed canopies were contrasted to assess the dynamic relationship between foliage adaptability to shading, stand LAI and tree growth. We found that individual leaf size is far more conservative in spruce than in beech. Individual leaf and needle area was larger at the top than at the bottom of the canopy in both species. Inverse relationship was found for specific leaf area (SLA), highest SLA values were found at lowest light availability under the canopy. There was no difference in leaf area index (LAI) between the two stands, however LAI increased from 10.8 to 14.6 m2m-2 between 2009 and 2011. Dominant trees of both species were more efficient in converting foliage mass or area to produce stem biomass, although this relationship changed with age and was species-specific. Overall, we found larger foliage plasticity in beech than in spruce in relation to light conditions, indicating larger capacity to exploit niche openings.

Does fertilizer type and method of application cause significant differences in essential oil yield and composition in rosemary (Rosmarinus officinalis L.)?

Organic fertilizers based on seaweed extract potentially have beneficial effects on many crop plants. Herewe investigate the impact of organic fertilizer on Rosmarinus officinalis measured by both yield and oilquality. Plants grown in a temperature-controlled greenhouse with a natural photoperiod and a controlledirrigation system were treated with seaweed fertilizer and an inorganic fertilizer of matching mineralcomposition but with no organic content. Treatments were either by spraying on to the foliage or wateringdirect to the compost. The essential oil was extracted by hydro-distillation with a Clevenger apparatusand analysed by gas-chromatography mass-spectrometry (GC–MS) and NMR. The chemical composi-tions of the plants were compared, and qualitative differences were found between fertilizer treatmentsand application methods. Thus sprayed seaweed fertilizer showed a significantly higher percentage of�-pinene, �-phellandrene, �-terpinene (monoterpenes) and 3-methylenecycloheptene than other treat-ments. Italicene, �-bisabolol (sesquiterpenes), �-thujene, and E-isocitral (monoterpenes) occurred insignificantly higher percentages for plants watered with the seaweed extract. Each was significantly dif-ferent to the inorganic fertilizer and to controls. The seaweed treatments caused a significant increasein oil amount and leaf area as compared with both inorganic treatments and the control regardless ofapplication method.

The biogeophysical climatic impacts of anthropogenic land use change during the Holocene

The first agricultural societies were established around 10 ka BP and had spread across much of Europe and southern Asia by 5.5 ka BP with resultant anthropogenic deforestation for crop and pasture land. Various studies (e.g. Joos et al., 2004; Kaplan et al., 2011; Mitchell et al., 2013) have attempted to assess the biogeochemical implications for Holocene climate in terms of increased carbon dioxide and methane emissions. However, less work has been done to examine the biogeophysical impacts of this early land use change. In this study, global climate model simulations with Hadley Centre Coupled Model version 3 (HadCM3) were used to examine the biogeophysical effects of Holocene land cover change on climate, both globally and regionally, from the early Holocene (8 ka BP) to the early industrial era (1850 CE). Two experiments were performed with alternative descriptions of past vegetation: (i) one in which potential natural vegetation was simulated by Top-down Representation of Interactive Foliage and Flora Including Dynamics (TRIFFID) but without land use changes and (ii) one where the anthropogenic land use model Kaplan and Krumhardt 2010 (KK10; Kaplan et al., 2009, 2011) was used to set the HadCM3 crop regions. Snapshot simulations were run at 1000-year intervals to examine when the first signature of anthropogenic climate change can be detected both regionally, in the areas of land use change, and globally. Results from our model simulations indicate that in regions of early land disturbance such as Europe and south-east Asia detectable temperature changes, outside the normal range of variability, are encountered in the model as early as 7 ka BP in the June–July–August (JJA) season and throughout the entire annual cycle by 2–3 ka BP. Areas outside the regions of land disturbance are also affected, with virtually the whole globe experiencing significant temperature changes (predominantly cooling) by the early industrial period. The global annual mean temperature anomalies found in our single model simulations were −0.22 at 1850 CE, −0.11 at 2 ka BP, and −0.03 °C at 7 ka BP. Regionally, the largest temperature changes were in Europe with anomalies of −0.83 at 1850 CE, −0.58 at 2 ka BP, and −0.24 °C at 7 ka BP. Large-scale precipitation features such as the Indian monsoon, the Intertropical Convergence Zone (ITCZ), and the North Atlantic storm track are also impacted by local land use and remote teleconnections. We investigated how advection by surface winds, mean sea level pressure (MSLP) anomalies, and tropospheric stationary wave train disturbances in the mid- to high latitudes led to remote teleconnections.