Δ13C and tree-ring width reflect different drought responses in Quercus ilex and Pinus halepensis

Holm oak (Quercus ilex L.) and Aleppo pine (Pinus halepensis Mill) are representative of two different functional types of trees extensively found in the Mediterranean: evergreen sclerophyllous and drought-adapted conifers. The former is considered a partially drought-tolerant species, whereas the latter is a typically drought-avoiding, water-saving species. We postulated that contrasting strategies in response to water deficits in Q. ilex and P. halepensis would lead to a differential sensitivity to changes in water availability. To test this hypothesis, we compared the response of both species in growth rate (measured as radial increments) and intrinsic water use efficiency [WUE_i, as inferred from carbon isotope discrimination (¹³C) in wood samples] among sites from different provenance regions in NE Spain. We found significant differences in ¹³C and growth among provenance regions, partly explained by contrasting water availability. Wood ¹³C was positively related with precipitation and the ratio between precipitation and potential evapotranspiration (P / E). However, these relationships were stronger in P. halepensis (for P / E, r ²=0.67, P <0.001) than in Q. ilex (r ²=0.42, P <0.01). In addition, radial growth was positively related with precipitation and ¹³C in P. halepensis (r ²=0.32 and r ²=0.35, respectively, P <0.01), but not in Q. ilex. We concluded that P. halepensis was more sensitive than Q. ilex to water availability, showing faster increase in WUE_i in response to water stress. We also found that the effect of north/south aspect on ¹³C and growth was site-specific, and unrelated to climatic variables.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Crop water availability in early agriculture: evidence from carbon isotope discrimination of seeds from a tenth millennium BP site on the Euphrates

The analysis of carbon isotope discrimination (Δ) in crop plant remains from archaeological sites may help to assess water availability for early agriculture. This study presents the analysis of Δ in seeds of naked wheat (Triticum aestivum/durum), lentil (Lens orientalis/culinaris), and flax (Linum sp.) found at the archaeological site of Tell Halula in the valley of the Middle Euphrates (Syria). This Neolithic site is the oldest in this region of the Fertile Crescent where the cultivation of domesticated plants has been reported, with seed remains ranging from 9550 to 8465 BP. Most of the seeds analysed showed Δ values greater than 16 ‰, reaching 20 ‰ for some samples of flax. For wheat, Δ values were much higher than those reported in present-day (1996) durum wheat crops cultivated under rainfed conditions in north-west Syria under environments with somewhat higher rainfall than Tell Halula. Similarly, grains of present-day (1997) barley cultivated in the archaeological site also showed lower values than those found in archaeological kernels. An empirical relationship between Δ of mature kernels and total precipitation (plus irrigation where applicable) from heading to maturity (r² = 0.82, n = 11) was established for durum wheat, currently cultivated in different environments of the Mediterranean basin. The resulting relationship was applied to the data on Δ of wheat fossil kernels from Tell Halula to estimate the accumulated water inputs during the time (about 6 weeks) the kernels were produced. Calculated water inputs for wheat during early agriculture were (over 110 mm) at least 5 times higher than current-day rainfall accumulated in Tell Halula during the same phenological period. These results strongly suggest that early agriculture wheat was cultivated at Tell Halula under much wetter conditions than are currently to be found in the area. The presence of flax and its very high Δ values also support this conclusion. Whether such humid conditions during cultivation were due to moister conditions prevailing at this time, by planting in alluvial areas or by irrigation works is discussed.     

Performance of hybrid poplar clones in short rotation coppice in Mediterranean environments: analysis of genotypic stability

Improving production in short rotation coppice (SRC) plantations requires, among other elements, a proper understanding of clonal performance. Genotypic stability over a range of environments is a factor of concern for breeding and recommendation purposes. Most common stability measures can be embedded in a mixed‐model framework accounting for interaction and heterocedasticity in genotype‐by‐environment tables. Data from nine hybrid poplars of different taxonomic background were tested in four Mediterranean sites under three agronomic practices (control, herbicide application, and supplementary fertilization) for total biomass (TB), stem biomass (SB), and branch biomass (BB) at the end of the first rotation. Stability models (stability variance, Finlay–Wilkinson and Eberhart–Russell) were compared, also allowing for the definition of groups of genotypes with distinct taxonomic backgrounds and a priori different variabilities. Results showed that genotype‐by‐environment (GE) interactions were associated with factors inherent to evaluation sites rather than to the agronomic practices tested. Depending on biomass fraction, regression models provided appropriate stability measures. Highly reactive clones to improving environmental conditions (e.g., ‘AF2’) tended to show the largest mean TB. However, this was not always the case, as clone ‘Monviso’ showed both intermediate reactivity (i.e., stable sensu Eberhart–Russell) and enhanced overall performance. The taxonomic group was relevant for explaining stability patterns for SB. The stability assessment for BB indicated different patterns in biomass allocation. Present findings point to the feasibility of either exploiting specific adaptation (in which case hybrid type may play a relevant role) or searching for broadly adapted, stable material exhibiting good performance in Mediterranean conditions.     

Climate-related variability in carbon and oxygen stable isotopes among populations of Aleppo pine grown in common-garden tests

The Aleppo pine (Pinus halepensis Mill.) is found in the Mediterranean under a broad range of moisture and thermal conditions. Differences in severity and duration of water stress among native habitats may act as selective forces shaping the populations’ genetic make-up in terms of contrasting drought strategies. We hypothesised that these strategies should translate into intraspecific variation in carbon isotope composition (δ¹³C, surrogate of intrinsic water-use efficiency, WUE_i) of wood holocellulose, and such variation might be linked to changes in oxygen isotope composition (δ¹⁸O, proxy of stomatal conductance) and to some climatic features at origin. Thus, we evaluated δ¹³C, δ¹⁸O, growth and survival for 25 Aleppo pine populations covering its geographic range and grown in two common-garden tests. We found intraspecific variability for δ¹³C and growth, with high-WUE_i populations (which showed ¹⁸O-enriched holocellulose) having low growth. These results suggest stomatal regulation as common control for δ¹³C and productivity. We also detected sizeable relationships between δ¹³C and climate factors related to the magnitude and timing of drought such as the ratio of summer to annual rainfall. The main climate variable associated with δ¹⁸O was minimum temperature, but only in the coldest trial, suggesting differences in growth rhythms among sources. Overall, slow growing populations from highly-seasonal dry areas of the western Mediterranean exhibited a conservative water-use, as opposed to fast growing sources from the northernmost distribution range. The particular behaviour of the Mediterranean Aleppo pine as compared with other conifers demonstrates different selective roles of climate variables in determining intraspecific fitness.     

Ground-Penetrating Radar as phenotyping tool for characterizing intraspecific variability in root traits of a widespread conifer

Plant and Soil - Drought is the main abiotic stress affecting Mediterranean forests. Root systems are responsible for water uptake, but intraspecific variability in tree root morphology is poorly...     

Climate at the onset of western Mediterranean agriculture expansion: Evidence from stable isotopes of sub-fossil oak tree rings in Spain

Climate conditions during the early Holocene may have contributed to the spread of Old World agriculture from its area of birth, in the Fertile Crescent (Near East), west through the Mediterranean Basin. Reconstructing the specific environmental conditions existing in early agricultural settlements of the western Mediterranean may help to elucidate this point. The aim of this work was to gain information on past climate of one of the earliest agricultural settlements of the Iberian Peninsula, La Draga, a lacustrine site dating back to the 2nd half of the 8th millennium BP, in which post fragments of deciduous oak have been exceptionally well preserved in an anaerobic environment (sub-fossil wood). We studied the relationship between climate factors and carbon isotope discrimination (Δ¹³C) and oxygen isotope composition (δ¹⁸O) in wood α-cellulose from modern tree-ring series of Quercus humilis Mill. and Quercus petraea (Matt) Liebl. recovered from the area. Climatic responses observed in extant material were used to interpret the isotopic signal of wood α-cellulose extracted from sub-fossil wood. Results showed that water availability of late spring, early summer and September influences Δ¹³C and δ¹⁸O, which allows their use as proxies for palaeoclimatic reconstruction. Differences between sub-fossil and extant samples in Δ¹³C (19.35 vs. 18.02‰) and δ¹⁸O (26.32 vs. 29.28‰) records suggest slightly lower temperatures and higher plant water availability than at present during the establishment of agriculture at the site. These results seem to disprove the existence of an environmental limitation preventing continuation of hunting and gathering activities as a cause for the adoption of agriculture in this early agricultural site on the Iberian Peninsula.     

Combined use of δ¹³C, δ18O and δ15N tracks nitrogen metabolism and genotypic adaptation of durum wheat to salinity and water deficit

? Accurate phenotyping remains a bottleneck in breeding for salinity and drought resistance. Here the combined use of stable isotope compositions of carbon (δ13C), oxygen (δ1?O) and nitrogen (δ1?N) in dry matter is aimed at assessing genotypic responses of durum wheat under different combinations of these stresses. ? Two tolerant and two susceptible genotypes to salinity were grown under five combinations of salinity and irrigation regimes. Plant biomass, δ13C, δ1?O and δ1?N, gas-exchange parameters, ion and N concentrations, and nitrate reductase (NR) and glutamine synthetase (GS) activities were measured. ? Stresses significantly affected all traits studied. However, only δ13C, δ1?O, δ1?N, GS and NR activities, and N concentration allowed for clear differentiation between tolerant and susceptible genotypes. Further, a conceptual model explaining differences in biomass based on such traits was developed for each growing condition. ? Differences in acclimation responses among durum wheat genotypes under different stress treatments were associated with δ13C. However, except for the most severe stress, δ13C did not have a direct (negative) relationship to biomass, being mediated through factors affecting δ1?O or N metabolism. Based upon these results, the key role of N metabolism in durum wheat adaptation to salinity and water stress is highlighted.     

Hydraulic and photosynthetic limitations prevail over root non-structural carbohydrate reserves as drivers of resprouting in two Mediterranean oaks

Resprouting is an ancestral trait in angiosperms that confers resilience after perturbations. As climate change increases stress, resprouting vigor is declining in many forest regions, but the underlying mechanism is poorly understood. Resprouting in woody plants is thought to be primarily limited by the availability of non-structural carbohydrate reserves (NSC), but hydraulic limitations could also be important. We conducted a multifactorial experiment with two levels of light (ambient, 2–3% of ambient) and three levels of water stress (0, 50 and 80 percent losses of hydraulic conductivity, PLC) on two Mediterranean oaks (Quercus ilex and Q. faginea) under a rain-out shelter (n = 360). The proportion of resprouting individuals after canopy clipping declined markedly as PLC increased for both species. NSC concentrations affected the response of Q. ilex, the species with higher leaf construction costs, and its effect depended on the PLC. The growth of resprouting individuals was largely dependent on photosynthetic rates for both species, while stored NSC availability and hydraulic limitations played minor and non-significant roles, respectively. Contrary to conventional wisdom, our results indicate that resprouting in oaks may be primarily driven by complex interactions between hydraulics and carbon sources, whereas stored NSC play a significant but secondary role.