Growth duration is a better predictor of stem increment than carbon supply in a Mediterranean oak forest: implications for assessing forest productivity under climate change.

Understanding whether tree growth is limited by carbon gain (source limitation) or by the direct effect of environmental factors such as water deficit or temperature (sink limitation) is crucial for improving projections of the effects of climate change on forest productivity. We studied the relationships between tree basal area (BA) variations, eddy covariance carbon fluxes, predawn water potential (Ψpd ) and temperature at different timescales using an 8-yr dataset and a rainfall exclusion experiment in a Quercus ilex Mediterranean coppice. At the daily timescale, during periods of low temperature (< 5°C) and high water deficit (< -1.1 MPa), gross primary productivity and net ecosystem productivity remained positive whereas the stem increment was nil. Thus, stem increment appeared limited by drought and temperature rather than by carbon input. Annual growth was accurately predicted by the duration of BA increment during spring (Δtt0-t1 ). The onset of growth (t0 ) was related to winter temperatures and the summer interruption of growth (t1 ) to a threshold Ψpd value of -1.1 MPa. We suggest that using environmental drivers (i.e. drought and temperature) to predict stem growth phenology can contribute to an improvement in vegetation models and may change the current projections of Mediterranean forest productivity under climate change scenarios.

Morphological and phenological shoot plasticity in a Mediterranean evergreen oak facing long-term increased drought.

Mediterranean trees must adjust their canopy leaf area to the unpredictable timing and severity of summer drought. The impact of increased drought on the canopy dynamics of the evergreen Quercus ilex was studied by measuring shoot growth, leaf production, litterfall, leafing phenology and leaf demography in a mature forest stand submitted to partial throughfall exclusion for 7 years. The leaf area index rapidly declined in the throughfall-exclusion plot and was 19% lower than in the control plot after 7 years of treatment. Consequently, leaf litterfall was significantly lower in the dry treatment. Such a decline in leaf area occurred through a change in branch allometry with a decreased number of ramifications produced and a reduction of the leaf area supported per unit sapwood area of the shoot (LA/SA). The leafing phenology was slightly delayed and the median leaf life span was slightly longer in the dry treatment. The canopy dynamics in both treatments were driven by water availability with a 1-year lag: leaf shedding and production were reduced following dry years; in contrast, leaf turnover was increased following wet years. The drought-induced decrease in leaf area, resulting from both plasticity in shoot development and slower leaf turnover, appeared to be a hydraulic adjustment to limit canopy transpiration and maintain leaf-specific hydraulic conductivity under drier conditions.

Amplified Drought Alters Leaf Litter Metabolome, Slows Down Litter Decomposition, and Modifies Home Field (Dis)Advantage in Three Mediterranean Forests.

In Mediterranean ecosystems, the projected rainfall reduction of up to 30% may alter plant-soil interactions, particularly litter decomposition and Home Field Advantage (HFA). We set up a litter transplant experiment in the three main forests encountered in the northern part of the Medi-terranean Basin (dominated by either Quercus ilex, Quercus pubescens, or Pinus halepensis) equipped with a rain exclusion device, allowing an increase in drought either throughout the year or concentrated in spring and summer. Senescent leaves and needles were collected under two precipitation treatments (natural and amplified drought plots) at their "home" forest and were left to decompose in the forest of origin and in other forests under both drought conditions. MS-based metabolomic analysis of litter extracts combined with multivariate data analysis enabled us to detect modifications in the composition of litter specialized metabolites, following amplified drought treatment. Amplified drought altered litter quality and metabolomes, directly slowed down litter decomposition, and induced a loss of home field (dis)advantage. No indirect effect mediated by a change in litter quality on decomposition was observed. These results may suggest major alterations of plant-soil interactions in Mediterranean forests under amplified drought conditions.

Fungal root symbionts and their relationship with fine root proportion in native plants from the Bolivian Andean highlands above 3,700 m elevation.

Here, we examined the colonization by fungal root symbionts in the cultivated Andean grain Chenopodium quinoa and in 12 species that dominate plant communities in the Bolivian Altiplano above 3,700 m elevation and explore for the possible relationships between fungal colonization and fine root proportion. The 12 most abundant species in the study area were consistently colonized by AMF and DSE. In contrast, the annual Andean grain C. quinoa showed negligible or absence of mycorrhizal fungi colonizing roots. On the other hand, C. quinoa, Junelia seriphioides and Chersodoma jodopappa were infected to a varying degree by the root pathogen Olpidium sp. We observed no relationship between AMF and DSE colonization and proportion of fine roots in the root system, but instead, the ratio between DSE and AMF colonization (ratio DSE/AMF) negatively related with proportion of fine roots. Our findings support the hypothesis regarding the importance of DSE at high altitudes and suggest a functional relationship between the rate of DSE/AMF and proportion of fine roots. The colonization by the root pathogen Olpidium sp. in C. quinoa deserves further study since this Andean grain is increasingly important for the local economy in these marginal areas.

Leaf traits capture the effects of land use changes and climate on litter decomposability of grasslands across Europe.

Land use and climate changes induce shifts in plant functional diversity and community structure, thereby modifying ecosystem processes. This is particularly true for litter decomposition, an essential process in the biogeochemical cycles of carbon and nutrients. In this study, we asked whether changes in functional traits of living leaves in response to changes in land use and climate were related to rates of litter potential decomposition, hereafter denoted litter decomposability, across a range of 10 contrasting sites. To disentangle the different control factors on litter decomposition, we conducted a microcosm experiment to determine the decomposability under standard conditions of litters collected in herbaceous communities from Europe and Israel. We tested how environmental factors (disturbance and climate) affected functional traits of living leaves and how these traits then modified litter quality and subsequent litter decomposability. Litter decomposability appeared proximately linked to initial litter quality, with particularly clear negative correlations with lignin-dependent indices (litter lignin concentr tion, lignin:nitrogen ratio, and fiber component). Litter quality was directly related to community-weighted mean traits. Lignin-dependent indices of litter quality were positively correlated with community-weighted mean leaf dry matter content (LDMC), and negatively correlated with community-weighted mean leaf nitrogen concentration (LNC). Consequently, litter decomposability was correlated negatively with community-weighted mean LDMC, and positively with community-weighted mean LNC. Environmental factors (disturbance and climate) influenced community-weighted mean traits. Plant communities experiencing less frequent or less intense disturbance exhibited higher community-weighted mean LDMC, and therefore higher litter lignin content and slower litter decomposability. LDMC therefore appears as a powerful marker of both changes in land use and of the pace of nutrient cycling across 10 contrasting sites.

Seasonal carbon storage and growth in Mediterranean tree seedlings under different water conditions.

In all Mediterranean-type ecosystems, evergreen and deciduous trees differing in wood anatomy, growth pattern and leaf habit coexist, suggesting distinct adaptative responses to environmental constraints. This study examined the effects of summer water stress on carbon (C) storage and growth in seedlings of three coexisting Mediterranean trees that differed in phenology and wood anatomy characteristics: Quercus ilex subsp. ballota (Desf.) Samp., Quercus faginea Lam. and Pinus halepensis L. Seedlings were subjected to two levels of watering during two consecutive summers and achieved a minimum of -0.5 and -2.5 MPa of predawn water potential in the control and water stress treatment, respectively. Both Quercus species concentrated their growth in the early growing season, demanding higher C in early spring but replenishing C-stores in autumn. These species allocated more biomass to roots, having larger belowground starch and lipid reserves. Quercus species differed in seasonal storage dynamics from P. halepensis. This species allocated most of its C to aboveground growth, which occurred gradually during the growing season, leading to fewer C-reserves. Soluble sugar and starch concentrations sharply declined in August in P. halepensis, probably because reserves support respiration demands as this species closed stomata earlier under water stress. Drought reduced growth of the three species, mainly in Q. faginea and P. halepensis, but not C-reserves, suggesting that growth under water stress conditions is not limited by C-availability.

Do elevation of CO(2) concentration and nitrogen fertilization alter storage and remobilization of carbon and nitrogen in pedunculate oak saplings?

Soil nitrogen can alter storage and remobilization of carbon and nitrogen in forest trees and affect growth responses to elevated carbon dioxide concentration ([CO(2)]). We investigated these effects in oak saplings (Quercus robur L.) exposed for two years to ambient or twice ambient [CO(2)] in combination with low- (LN, 0.6 mmol N l(-1)) or high-nitrogen (HN, 6.1 mmol N l(-1)) fertilization. Autumn N retranslocation efficiency from senescing leaves was less in HN saplings than in LN saplings, but about 15% of sapling N was lost to the litter. During the dormant season, nonstructural carbohydrates made up 20 to 30% of the dry mass of perennial organs. Starch was stored mainly in large roots where it represented 35-46% of dry mass. Accumulation of starch increased in large roots in response to LN but was unaffected by elevated [CO(2)]. The HN treatment resulted in high concentrations of N-soluble compounds, and this effect was reduced by elevated [CO(2)], which decreased soluble protein N (-17%) and amino acid N (-37%) concentrations in the HN saplings. Carbon and N reserves were labeled with (13)C and (15)N, respectively, at the end of the first year. In the second year, about 20% of labeled C and 50% of labeled N was remobilized for spring growth in all treatments. At the end of leaf expansion, 50-60% of C in HN saplings originated from assimilation versus only 10-20% in LN saplings. In HN saplings only, N uptake occurred, and some newly assimilated N was allocated to new shoots. Through effects on the C and N content of perennial organs, elevated [CO(2)] and HN increased remobilization capacity, thereby supporting multiple shoot flushes, which increased leaf area and subsequent C acquisition in a positive feedback loop.

Efficiency of near-infrared reflectance spectroscopy to assess and predict the stage of transformation of organic matter in the composting process.

Changes in composts of sewage sludges and green wastes were analysed by near infrared reflectance spectroscopy (NIRS) and chemical analysis with 426 samples representative of six stages of composting: 8, 20, 35, 75, 135 and 180 days. Maturity of compost was assessed through changes in C:N ratio. Results of spectroscopic properties (200 wavelengths) were studied with several multivariate analyses. First, a descriptive approach revealed compost changes with time of maturation. Then, a constrained ordination (RDA with permutation tests) demonstrated a significant effect of three factors of sampling: stage of composting, depth and position in windrows. Precise calibration models between spectral data, the C, N, C:N values and composting time were build using partial least square regression (r(2)>0.95). Together, these results show the efficiency of NIRS to predict chemical changes and the stage of transformation of organic matter during the composting process.

Discontinuities in quinoa biodiversity in the dry Andes: An 18-century perspective based on allelic genotyping.

History and environment shape crop biodiversity, particularly in areas with vulnerable human communities and ecosystems. Tracing crop biodiversity over time helps understand how rural societies cope with anthropogenic or climatic changes. Exceptionally well preserved ancient DNA of quinoa (Chenopodium quinoa Willd.) from the cold and arid Andes of Argentina has allowed us to track changes and continuities in quinoa diversity over 18 centuries, by coupling genotyping of 157 ancient and modern seeds by 24 SSR markers with cluster and coalescence analyses. Cluster analyses revealed clear population patterns separating modern and ancient quinoas. Coalescence-based analyses revealed that genetic drift within a single population cannot explain genetic differentiation among ancient and modern quinoas. The hypothesis of a genetic bottleneck related to the Spanish Conquest also does not seem to apply at a local scale. Instead, the most likely scenario is the replacement of preexisting quinoa gene pools with new ones of lower genetic diversity. This process occurred at least twice in the last 18 centuries: first, between the 6th and 12th centuries-a time of agricultural intensification well before the Inka and Spanish conquests-and then between the 13th century and today-a period marked by farming marginalization in the late 19th century likely due to a severe multidecadal drought. While these processes of local gene pool replacement do not imply losses of genetic diversity at the metapopulation scale, they support the view that gene pool replacement linked to social and environmental changes can result from opposite agricultural trajectories.

Rain-fed agriculture thrived despite climate degradation in the pre-Hispanic arid Andes.

Archaeological research suggests significant human occupation in the arid Andean highlands during the 13th to 15th centuries, whereas paleoclimatic studies reveal prolonged drier and colder conditions during that period. Which subsistence strategy supported local societies in this harsh environment? Our field and aerial surveys of archaeological dwelling sites, granaries, and croplands provide the first evidence of extended pre-Hispanic agriculture supporting dense human populations in the arid Andes of Bolivia. This unique agricultural system associated with quinoa cultivation was unirrigated, consisting of simple yet extensive landscape modifications. It relied on highly specific environmental knowledge and a set of water-saving practices, including microterracing and biennial fallowing. This intense agricultural activity developed during a period of unfavorable climatic change on a regional and global scale, illustrative of efficient adaptive strategies to cope with this climatic change.

Interspecific relationships among soil invertebrates influence pollutant effects of phenanthrene.

Five mesofauna communities varying in both structure and composition were exposed to phenanthrene in mesocosms for up to four months. Effects of phenanthrene were assessed on mesofauna population dynamics, fungal biomass (ergosterol concentrations), and litter decomposition (litter mass loss, nitrogen concentration). The effects of each community on the fate of phenanthrene were also assessed. We hypothesize that phenanthrene affects the population dynamics of mesofauna and soil biological functioning depending on exposure duration, type of community, or both. Results show that phenanthrene exerted an effect on mesofauna and that the effects on some species, like Folsomia fimetaria, were influenced by the species composition in the mesocosms, the soil layer, and the sampling date. However, the effects of phenanthrene on ergosterol content and organic matter decomposition were not significantly influenced by community composition. These results demonstrate that interspecific relationships are needed to assess the toxicity of pollutants and should be taken into account in ecotoxicological risk assessment. Furthermore, this work does not support the hypothesis of a direct link between toxic effects of organic pollutants on mesofauna species and soil biological functioning.

Sewage sludge effects on mesofauna and cork oak (Quercus suber L.) leaves decomposition in a Mediterranean forest firebreak.

Effects of sewage sludge on litter mesofauna communities (Collembola and Acari) and cork oak (Quercus suber L.) leaf litter decomposition have been studied during 18 mo using litterbags in an in situ experimental forest firebreak in southeastern France. The sludge (2.74 t DM ha(-1) yr(-1)) was applied to fertilize and maintain a pasture created on the firebreak. Litterbag colonization had similar dynamics on both the control and fertilized plots and followed a typical Mediterranean pattern showing a greater abundance in spring and autumn and a lower abundance in summer. After 9 mo of litter colonization, Collembola and Acari, but mainly Oribatida, were more abundant on the sludge-fertilized plot. Leaf litter decomposition showed a similar pattern on both plots, but it was faster on the control plot. Furthermore, leaves from the fertilized plot were characterized by greater nitrogen content. Both chemical composition of leaves and sludges and the decomposition state of leaves have significantly affected the mesofauna community composition from each plot.

Isoprenoid emissions of Quercus spp. (Q. suber and Q. ilex) in mixed stands contrasting in interspecific genetic introgression.

• Among oak species, Quercus ilex is classified as a monoterpene emitter and Q. suber is mainly known as a nonisoprenoid emitter. The extent and origin of this diversification is unknown. • We examined intra- and interspecific emission variability in two mixed stands which differed in their level of hybridization and reciprocal genetic introgression based on variations in cytoplasmic (chloroplast DNA) and nuclear (allozyme) markers. • At both sites all trees identified as Q. ilex, or as recent descendants from Q. ilex × Q. suber hybrids, emitted monoterpenes. Of Q. suber trees (genetically introgressed or not by Q. ilex), 91% were also monoterpene emitters, and the remainder nonemitters. One tree identified as a Q. canariensis × Q. ilex hybrid emitted both isoprene and monoterpenes. Compared with Q. ilex, the standard emission rate of Q. suber was higher in summer and lower in autumn. Both species emitted the same monoterpenes, proportions of which showed significant intra- and interspecific variability. • The results suggest that Q. suber populations in the French Mediterranean intrinsically emit monoterpenes, and that gene flow between oak species contributes to diversification of emission signatures.

Using near-infrared reflectance spectroscopy to predict carbon, nitrogen and phosphorus content in heterogeneous plant material.

The aim of this study was to produce calibration equations between near-infrared reflectance (NIR) spectra and the concentrations of carbon, nitrogen, and phosphorus in heterogeneous material: from living needles to litter in Pinus halepensis stands subjected to prescribed burnings. The aim was to determine whether calibrations should be conducted within each stage in the transformation of needles (local calibrations), giving relationships that were accurate but valid only for each particular stage, or whether it was possible to integrate the various forms of variation in needles (global calibrations) while retaining an acceptable accuracy. A principal component analysis calculated from the sample spectral data was used to distinguish three different sets, each sharing spectral characteristics and corresponding to three categories of needle: needles collected on the pines (N), falling needles (F), and litter (L), and each containing samples collected from the burnt sites and a control site. Samples representative of all the forms of variation in spectral properties were selected from within each category and their carbon, nitrogen, and phosphorus concentrations were measured using standard wet chemistry methods; these constituted the calibration sets n, f, and l. Calibrations were produced between the nutrient concentrations and the NIR spectra of the calibration sets n, f, and l and the grouped sets (n+f, f+l, n+f+l). The results of local calibrations made from each individual category showed that the carbon, nitrogen, and phosphorus concentrations were accurately predictable by NIR spectra. The global calibrations made by lumping together several categories were valid for a wider range of concentrations and for spectrally heterogeneous materials and in most cases were just as accurate as the local calibrations produced from each individual category.

Calibration of chemical and biological changes in cocomposting of biowastes using near-infrared spectroscopy.

Cocomposting of green wastes and sewage sludges is a complex process involving rapid biological and chemical changes. The objective of the study was to assess the usefulness of near-infrared reflectance spectroscopy (NIRS) to characterize these changes, as an alternative to standard procedures which are often time-consuming and laborious. Samples obtained during 146 days of composting were analyzed by 14 conventional methods and NIRS. Results from conventional methods demonstrated a noticeable separation into two distinct phases. An initial phase from 4 to 50-60 days was characterized by intensive degradation. A second phase up to 146 days was characterized by a decrease in all biological activities. NIRS calibrations allowed accurate predictions of nitrogen (N), carbon (C), C/N, humic acid (HA), pH, respiration, cellulase, phenoloxidase, and composting time successfully. Results were less accurate for organic matter (OM), protease, acid, and alkaline phosphatases and unsatisfactory for fulvic acid. NIRS calibration allows composting time/state of progress of maturation to be predicted accurately to within 10 days. A global index of composting evolution (GICE), resulting from the 14 parameters studied, is proposed. It is precisely predicted and shows that since NIRS is able to predict essential parameters of compost maturity, it could prove invaluable for monitoring biowastes cocomposting.

Quantifying species composition in root mixtures using two methods: near-infrared reflectance spectroscopy and plant wax markers.

Understanding of plant interactions is greatly limited by our ability to identify and quantify roots belonging to different species. We proposed and compared two methods for estimating the root biomass proportion of each species in artificial mixtures: near-infrared reflectance spectroscopy (NIRS) and plant wax markers. Two sets of artificial root mixtures composed of two or three herbaceous species were prepared. The proportion of root material of each species in mixtures was estimated from NIRS spectral data (i) and the concentration patterns of n-alkanes (ii), n-alcohols (iii), and n-alkanes +n-alcohols combined (iv). For each data set, calibration equations were developed using multivariate statistical models. The botanical composition of root mixtures was predicted well for all the species considered. The accuracy varied slightly among methods: alkanes < alcohols = alkanes + alcohols < NIRS. Correlation coefficients between predicted and actual root proportions ranged from 0.89 to 0.99 for alkanes + alcohols predictions and from 0.97 to 0.99 for NIRS predictions. These two methods provide promising potential for understanding allocation patterns and competitive interactions.

Leaf morphology, photochemistry and water status changes in resprouting Quercus ilex during drought.

Functional and morphological (structural) characteristics of Quercus ilex L. leaves under drought stress were studied in the forest and in a nursery. We compared undisturbed individuals (controls) with resprouts emerging after clear-cut or excision. When soil water availability was high, gas-exchange was similar in resprouts and controls, despite higher midday leaf water potential, midday leaf hydration and relative water content (RWC). In moderate drought, stomatal closure was found to limit photosynthesis in controls, and in severe drought non-stomatal limitations of photosynthesis were also greater than in resprouts. Leaf structure and chemical composition changed under drought stress. Leaves tended to be smaller in controls with increasing drought, and resprouts had larger leaves and lower leaf mass area (LMA). The relationship between nitrogen (N) content and LMA implied lower N investment in photosynthetic components in controls, which could be responsible for their increased non-stomatal limitation of photosynthesis. Changes were more apparent in leaf density (D) and thickness (T), components of LMA. Decreases in D were related to reductions in cell wall components: hemicellulose, cellulose and lignin. In resprouts, reduced D and leaf T accounted for the higher mesophyll conductance (gmes) to CO2 measured.

The function of the superficial root mat in the biogeochemical cycles of nutrients in congolese eucalyptus plantations.

BACKGROUND AND AIMS: The importance of superficial root mats inside the forest floor for the nutrition of Amazonian rain forests has been extensively investigated. The present study was aimed at assessing the function of a root mat adherent to decomposing organic material observed in Eucalyptus plantations. METHODS: The development of the root mat was studied through micromorphological observations of thin litter sections, and the influence of soil microtopography and soil water repellency on root mat biomass was assessed in situ on an area of 5 m2. In addition, input-output budgets of nutrients within the forest floor were established from measurements of litterfall, dissolved nutrients in gravitational solutions, and forest floor nutrient contents. KEY FINDINGS: The amounts of nutrients released during litter decay in this ecosystem during the period of study were, on average, 46, 3, 4, 19 and 17 kg ha-1 year-1 for N, P, K, Ca and Mg, respectively. The simultaneous measurements of the chemical composition of throughfall solutions and leachates beneath the forest floor showed a very quick uptake of nutrients by the root mat during the decomposition processes. Indeed, the solutions did not become noticeably enriched in nutrients during their passage through the holorganic layer, despite large amounts of elements being released during litter decay. The root mat biomass decreased significantly during the dry season, and a preferential development in microdepressions at the soil surface was observed. A strong water repellency observed in these depressions might enhance the ability of the roots to take up water and nutrients during the dry periods. CONCLUSIONS: The root mat was active throughout the year to catch the flux of nutrients from the biodegradation of the forest floor, preventing the transfer of dissolved nutrients toward deeper soil horizons. This mechanism is involved in the successful adaptation of this Eucalyptus hybrid in areas covered by 'climacic' savannas in Congo.

A generic model to describe the dynamics of nutrient concentrations within stemwood across an age series of a eucalyptus hybrid.

Nutrient concentrations (N, P, K) were determined within stemwood in an age series of eucalyptus stands. Four trees per stand were selected according to their size to represent the whole range of basal areas in 1-, 2-, 3-, 4-, 5-, 6- and 7-year-old stands. Cross-sections were sampled every 4 m from the ground to the top of the tree, and chemical analyses were performed for each annual ring in the cross-sections. We constructed a new and generic model to describe the dynamics of nutrient concentrations within the stemwood. Three main parameters were used: (1) the initial concentration of the ring, Ic; (2) the final concentration of the ring at harvest, Fc; and (3) the rate of change in concentration, k. The model is very flexible and was adapted to describe N, P and K concentrations within the stems, and their dynamics over time. An analysis of the parameters showed that k was constant for a given nutrient. Ic varied with height within the tree for P, whereas for N and K it was a function of: (1) the age of the tree when the ring was initiated: and (2) height within the tree. Fc was constant for N, and dependent on the age of the tree when the ring was initiated for K and P. The final models showed a low Root Mean Square Error for a limited number of parameters (less than seven). When validated on an independent sample, the models were shown to have high predictive quality.