Holocene changes in precipitation seasonality in the western Mediterranean Basin: a multi-species approach using d 13 C of archaeobotanical remains

ABSTRACT: Precipitation has been of utmost importance in shaping the evolution of landscapes and human settlements in the Mediterranean. However, information on seasonal precipitation patterns through the Holocene is scarce. This study attempts to quantify the evolution of seasonal precipitation in the East Iberian Peninsula (5000 BC to AD 600) based on the carbon isotope composition (d 13 C) of archaeobotanical remains. Data on Holm oak, Aleppo pine and small-grain cereals were combined, and precipitation was inferred from models relating present-day records to the d 13 C of modern samples. Subsequently, charred grains were used as a proxy for ancient moisture during April-May, whereas oak and pine charcoals provided complementary rainfall estimates for September-December and January-August, respectively. The results reveal aridity changes throughout the Holocene in the western Mediterranean. Past spring-summer precipitation was consistently higher than at present. In contrast, autumn and early winter precipitation showed stronger fluctuations, particularly during the first millennium BC, and often exhibited values below those of the present. The high contribution of autumn precipitation to the annual water budget, typical of the present Mediterranean climate, was definitively established at the beginning of the current era. This study shows how a combination of species holding complementary environmental signals can contribute to a wider knowledge of local precipitation dynamics

Increasing drought effects on five European pines modulate Δ13C-growth coupling along a Mediterranean altitudinal gradient.

Climate warming increases vulnerability to drought in Mediterranean water-limited forests. However, we still lack knowledge of the long-term physiological responses of coexisting pine species in these forests regarding their ability to cope with warming-induced drought stress. We investigated spatiotemporal patterns of tree performance for five isohydric pines with partially overlapping ecological niches in the eastern Iberian Peninsula along an altitudinal gradient: Pinus halepensis = P. pinaster ≤ P. nigra ≤ P. sylvestris ≤ P. uncinata. Using indexed tree-ring widths (TRWi) we assessed changes in temporal coherence of radial growth (growth synchrony, âC) over the period 1902-2011 across three elevation belts: low ≈ 1100 m; mid = 1615 m; high = 2020 m. We also examined by mixed modelling whether TRWi showed an increased coupling with leaf-level gas exchange (inferred from indexed carbon isotope discrimination, Δ13Ci) by enhanced stomatal regulation in response to an amplified regional drought stress. Increasingly negative annual water balances (decrease in annual precipitation minus evapotranspiration = -4.8 mm year−1; 1970-2011) prompted more synchronous growth of coexisting pines between low- and mid-elevation belts, with âC rising from 0.25 ± 0.04 (1902-1951) to 0.62 ± 0.05 (1962-2011). This effect was coupled with tighter stomatal regulation at mid-elevation as indicated by high correlations between TRWi and Δ13Ci (>0.60 from the mid-1970s onwards) which resembled those found at low-elevation. Simultaneously, TRWi vs. Δ13Ci uncoupling occurred at the high-elevation belt across species. Weaker growth-climate relationships as elevation increased highlighted the major role of the altitude-dependent thermal gradient in growth responsiveness to drought; however, an intensified Δ13Ci response to spring water availability across elevation belts observed from mid-1970s onwards suggested regional shifts in tree physiological activity linked to earlier seasonal drought impacts. Warming-induced drought stress is spreading to higher altitudes in Iberian pinewoods as multispecies growth is linked to progressively tighter stomatal control of water losses reflected in wood Δ13C

Carbon stable isotope analysis of cereal remains as a way to reconstruct water availability: preliminary results

Reconstructing past water availability, both as rainfall and irrigation, is important to answer questions about the way society reacts to climate and its changes and the role of irrigation in the development of social complexity. Carbon stable isotope analysis of archaeobotanical remains is a potentially valuable method for reconstructing water availability. To further define the relationship between water availability and plant carbon isotope composition and to set up baseline values for the Southern Levant, grains of experimentally grown barley and sorghum were studied. The cereal crops were grown at three stations under five different irrigation regimes in Jordan. Results indicate that a positive but weak relationship exists between irrigation regime and total water input of barley grains, but no relationship was found for sorghum. The relationship for barley is site-specific and inter-annual variation was present at Deir ‘Alla, but not at Ramtha and Khirbet as-Samra

High-carotenoid maize: development of plant biotechnology prototypes for human and animal health and nutrition

Carolight (R) is a transgenic maize variety that accumulates extraordinary levels of carotenoids, including those with vitamin A activity. The development of Carolight (R) maize involved the technical implementation of a novel combinatorial transformation method, followed by rigorous testing for transgene expression and the accumulation of different carotenoid molecules. Carolight (R) was envisaged as a way to improve the nutritional health of human populations that cannot access a diverse diet, but this ultimate humanitarian application can only be achieved after extensive testing for safety, agronomic performance and nutritional sufficiency. In this article, we chart the history of Carolight (R) maize focusing on its development, extensive field testing for agronomic performance and resistance to pests and pathogens, and feeding trials to analyze its impact on farm animals (and their meat/dairy products) as well as animal models of human diseases. We also describe more advanced versions of Carolight (R) endowed with pest-resistance traits, and other carotenoid-enhanced maize varieties originating from the same series of initial transformation experiments. Finally we discuss the further steps required before Carolight (R) can fulfil its humanitarian objectives, including the intellectual property and regulatory constraints that lie in its path

Mesophyll diffusion conductance to CO 2: An unappreciated central player in photosynthesis

Mesophyll diffusion conductance to CO 2 is a key photosynthetic trait that has been studied intensively in the past years. The intention of the present review is to update knowledge of g m, and highlight the important unknown and controversial aspects that require future work. The photosynthetic limitation imposed by mesophyll conductance is large, and under certain conditions can be the most significant photosynthetic limitation. New evidence shows that anatomical traits, such as cell wall thickness and chloroplast distribution are amongst the stronger determinants of mesophyll conductance, although rapid variations in response to environmental changes might be regulated by other factors such as aquaporin conductance.Gaps in knowledge that should be research priorities for the near future include: how different is mesophyll conductance among phylogenetically distant groups and how has it evolved? Can mesophyll conductance be uncoupled from regulation of the water path? What are the main drivers of mesophyll conductance? The need for mechanistic and phenomenological models of mesophyll conductance and its incorporation in process-based photosynthesis models is also highlighted.The study was financially supported by the Estonian Ministry of Science and Education (grant SF1090065s07), the Spanish Ministry of Science and Innovation through projects BFU2008-01072 (MEFORE), AGL2009-11310/AGR, BFU2011-23294 (MECOME) and CGL2009-13079-C02-01 (PALEOISOTREE), and the European Commission through European Regional Fund (the Estonian Center of Excellence in Environmental Adaptation), and the Marie Curie project MC-ERG-246725 (FP7). J.P.F. is supported by the Ramón y Cajal program (RYC-2008-02050). A.G. had a Swiss National Science Fellowship (PA00P3_126259). M.M.B. and C.R.W are supported by Future Fellowships from the Australian Research Council (FT0992063 and FT100100024). C.D. was supported by a grant from the French government and by the cooperation project Tranzfor (Transferring Research between EU and Australia–New Zealand on Forestry and Climate Change, PIRSES-GA-2008-230793) funded by the European Union

Stable carbon Isotope evidence for neolithic and bronze age crop water management in the eastern mediterranean and southwest asia

In a large study on early crop water management, stable carbon isotope discrimination was determined for 275 charred grain samples from nine archaeological sites, dating primarily to the Neolithic and Bronze Age, from the Eastern Mediterranean and Western Asia. This has revealed that wheat (Triticum spp.) was regularly grown in wetter conditions than barley (Hordeum sp.), indicating systematic preferential treatment of wheat that may reflect a cultural preference for wheat over barley. Isotopic analysis of pulse crops (Lens culinaris, Pisum sativum and Vicia ervilia) indicates cultivation in highly varied water conditions at some sites, possibly as a result of opportunistic watering practices. The results have also provided evidence for local land-use and changing agricultural practices

Diversity of a cytokinin dehydrogenase gene in wild and cultivated barley

The cytokinin dehydrogenase gene HvCKX2.1 is the regulatory target for the most abundant heterochromatic small RNAs in drought-stressed barley caryopses. We investigated the diversity of HvCKX2.1 in 228 barley landraces and 216 wild accessions and identified 14 haplotypes, five of these with ten or more members, coding for four different protein variants. The third largest haplotype was abundant in wild accessions (51 members), but absent from the landrace collection. Protein structure predictions indicated that the amino acid substitution specific to haplotype 3 could result in a change in the functional properties of the HvCKX2.1 protein. Haplotypes 1–3 have overlapping geographical distributions in the wild population, but the average rainfall amounts at the collection sites for haplotype 3 plants are significantly higher during November to February compared to the equivalent data for plants of haplotypes 1 and 2. We argue that the likelihood that haplotype 3 plants were excluded from landraces by sampling bias that occurred when the first wild barley plants were taken into cultivation is low, and that it is reasonable to suggest that plants with haplotype 3 are absent from the crop because these plants were less suited to the artificial conditions associated with cultivation. Although the cytokinin signalling pathway influences many aspects of plant development, the identified role of HvCKX2.1 in the drought response raises the possibility that the particular aspect of cultivation that mitigated against haplotype 3 relates in some way to water utilization. Our results therefore highlight the possibility that water utilization properties should be looked on as a possible component of the suite of physiological adaptations accompanying the domestication and subsequent evolution of cultivated barley

Relationships between δ13C, δ18O and grain yield in bread wheat genotypes under favourable irrigated and rain-fed conditions

In previous investigations, carbon isotope composition (δ13C) has been used in C3 cereals to screen for genotypes with high transpiration efficiency and oxygen isotope composition (δ18O) has been shown to correlate with transpiration rate. We examined associations of δ13C of the grain and flag leaf and δ18O of the flag leaf with respect to grain yield in wheat cultivars in UK field conditions. Field experiments were carried out at University of Nottingham in 2009–10 and 2010–11 testing 17 wheat cultivars under fully irrigated and rain-fed conditions. Averaging across years grain yield was reduced by 1.69 t ha−1 (16.5%) in the rain-fed treatment (P < 0.001). There was a negative linear relationship between grain yield and grain δ13C amongst cultivars, under both irrigated (R2 = 0.47, P < 0.01) and rain-fed (R2 = 0.70, P < 0.001) conditions. Grain δ13C was negatively correlated with flag-leaf stomatal conductance (r = −0.94, P < 0.01) in a subset of six of the cultivars, indicating that higher transpiration efficiency was associated with lower stomatal conductance. The associations between grain yield and flag-leaf δ13C and flag-leaf δ18O amongst cultivars under irrigated and rain-fed conditions were not statistically significant. There was a positive linear relationship between flag-leaf δ18O and grain δ13C amongst cultivars under irrigated conditions (R2 = 0.38, P < 0.01), indicating a trade-off between transpiration and transpiration efficiency (TE). Genetic variation in grain yield under rain-fed conditions was also associated with delayed onset of flag-leaf senescence (R2 = 0.35, P < 0.05). The 17 wheat cultivars ranged in year of release (YoR) from 1964 to 2009 and grain yield increased linearly under irrigated conditions by 60.4 kg ha−1 yr−1 (0.72% yr−1) and under rain-fed conditions by 47.5 kg ha−1 yr−1 (0.66% yr−1) over the 45 year period and grain δ13C composition decreased by 0.0255 and 0.0304‰ yr−1, respectively, indicating genetic gains in wheat yield potential in the UK seem likely to have been achieved through a lower TE, higher water uptake and lesser limitation of stomatal conductance

Circadian rhythms have significant effects on leaf-to-canopy scale gas exchange under field conditions

Background Molecular clocks drive oscillations in leaf photosynthesis, stomatal conductance, and other cell and leaf-level processes over ∼24 h under controlled laboratory conditions. The influence of such circadian regulation over whole-canopy fluxes remains uncertain; diurnal CO2 and H2O vapor flux dynamics in the field are currently interpreted as resulting almost exclusively from direct physiological responses to variations in light, temperature and other environmental factors. We tested whether circadian regulation would affect plant and canopy gas exchange at the Montpellier European Ecotron. Canopy and leaf-level fluxes were constantly monitored under field-like environmental conditions, and under constant environmental conditions (no variation in temperature, radiation, or other environmental cues). Results We show direct experimental evidence at canopy scales of the circadian regulation of daytime gas exchange: 20–79 % of the daily variation range in CO2 and H2O fluxes occurred under circadian entrainment in canopies of an annual herb (bean) and of a perennial shrub (cotton). We also observed that considering circadian regulation improved performance by 8–17 % in commonly used stomatal conductance models. Conclusions Our results show that circadian controls affect diurnal CO2 and H2O flux patterns in entire canopies in field-like conditions, and its consideration significantly improves model performance. Circadian controls act as a ‘memory’ of the past conditions experienced by the plant, which synchronizes metabolism across entire plant canopies