Wheat genotypic variability in grain yield and carbon isotope discrimination under Mediterranean conditions assessed by spectral reflectance

A collection of 368 advanced lines and cultivars of spring wheat（Triticum aestivum L.） from Chile, Uruguay, and CIMMYT（Centro Internacional de Mejoramiento de Maíz y Trigo）, with good agronomic characteristics were evaluated under the Mediterranean conditions of central Chile. Three different water regimes were assayed： severe water stress（SWS, rain fed）, mild water stress（MWS; one irrigation around booting）, and full irrigation（FI; four irrigations： at tillering,flag leaf appearance, heading, and middle grain filling）. Traits evaluated were grain yield（GY）, agronomical yield components,days from sowing to heading, carbon isotope discrimination（△^13C） in kernels, and canopy spectral reflectance. Correlation analyses were performed for 70 spectral reflectance indices（SRI） and the other traits evaluated in the three trials. GY and △^13C were the traits best correlated with SRI, particularly when these indices were measured during grain filling. However,only GY could be predicted using a single regression, with ResearchNormalized Difference Moisture Index（NDMI2： 2,200; 1,100）having the best fit to the data for the three trials. For △^13C, only individual regressions could be forecast under FI（r^2： 0.25–0.37）and MWS（r^2： 0.45–0.59） but not under SWS（r^2： 0.03–0.09）.NIR‐based SRI proved to be better predictors than those that combine visible and NIR wavelengths.     

Relationships between carbon isotope discrimination, dry matter production, and harvest index in durum wheat

Carbon isotope discrimination has been proposed as a criterion for the indirect selection to improve transpiration efficiency and grain yield in bread wheat and barley. Less attention has been devoted to durumwheat (Triticum durumDesf.) despite its economic importance in the Mediterranean basin. The Δ genetic variation and its relationships to dry matter production and harvest index in durum wheat were investigated in this study. For this purpose, field experiments were conducted on 144 durum wheat accessions under Mediterranean conditions (South of France) during three consecutive years with contrasting climatic conditions. Grain yield, above-ground biomass, harvest index, and carbon isotope discrimination of flag leaf and kernel were measured. Differences between years, noted for both leaf and kernel carbon isotope discrimination, were probably related to the variation in water availability from year to year. A large genotypic variation was also noticed for both leaf and kernel carbon isotope discrimination. The two traits were found to be positively correlated with grain yield within and across years, which confirms the interest in carbon isotope discrimination for selection for grain yield improvement under Mediterranean conditions. Both kernel and leaf carbon isotope discrimination correlated better with harvest index than with grain yield, suggesting that carbon isotope discrimination could reflect the efficiency of carbon partitioning to the kernel. The lack of correlation between leaf carbon isotope discrimination and both harvest index and grain yield in favourable water conditions (1996) was probably due to the difference in water availability between the period until flag leaves sampling (favourable conditions) and the strong water stress which accompanied the grain filling. Kernel carbon isotope discrimination correlated better with both harvest index and grain yield than did leaf carbon isotope discrimination. Moreover, a higher broad-sense heritability was obtained for kernel carbon isotope discrimination than for leaf carbon isotope discrimination. As a result, kernel carbon isotope discrimination appeared to be a better predictive criterion for efficiency of the carbon partitioning to the kernel (harvest index), and hence for grain yield, than did flag leaf carbon isotope discrimination.     

Growth,abscisic acid content,and carbon isotope composition in wheat cultivars grown under different soil moisture

Changes in dry matter accumulation and allocation, abscisic acid content and carbon isotope composition of three wheat cultivars from dry, middle and wet climate regions were recorded at full maturity after exposure to different watering regimes (100, 50 and 25 % field capacity). Compared with the wet climate cultivar, the dry climate cultivar showed lower stem height, total leaf area, total dry biomass and total grain dry mass, and higher root/shoot ratio, abscisic acid content and carbon isotope composition under all watering regimes. Both water-limited treatments significantly reduced leaf growth and increased dry matter allocation into the roots leading to a significant raise of root/shoot ratio in all cultivars tested. In addition, drought affected morphological and physiological properties more in the dry climate cultivar than in the wet climate cultivar.     

Relationships between flag leaf carbon isotope discrimination and several morpho-physiological traits in durum wheat genotypes under Mediterranean conditions

Relationships between flag leaf carbon isotope discrimination (Delta), water status parameters, residual transpiration (RT) and stomatal density (SD) were examined on a collection of 144 durum wheat accessions. Associations between Delta, grain yield (GY) and harvest index (HI) were also studied. The field trial was conducted under Mediterranean conditions. The crop cycle was characterised by a period of drought from February until maturity. A broad range of values we obtained for Delta (16.5-19.9 per thousand) and other physiological traits. Flag leaf Delta was positively and significantly correlated with both HI and GY. Delta was better correlated with HI than with GY, which suggests that higher Delta values indicate higher efficiency of carbon partitioning to the kernel, leading to higher GY. Delta was found positively related with RT and negatively related with SD. This relationship may indicate a possible SD component of RT due to the association between conductance and SD. Strong positive correlations were found between Delta and water status parameters, suggesting that Delta may provide a good indication of plant water status in durum wheat under rainfed Mediterranean conditions.     

Productivity, carbon isotope discrimination and leaf traits of trees of Eucalyptus globulus Labill. in relation to water availability

The stand basal area, carbon isotope discrimination (Δ) in tree rings and leaves, leaf area index and leaf traits of trees were measured in 6‐ to 8‐year‐old stands of Eucalyptus globulus Labill. across a gradient of rainfall of 600–1400 mm year^?1 in south‐western Australia to better understand the importance of leaf traits and gas‐exchange as determinants of stand productivity. Δ ranged from 17‰ to 21‰. Δ and basal area were highly, positively correlated with each other and the ratio of mean annual rainfall to potential evaporation (P/PE). Leaf area index, soil water holding capacity and leaf nitrogen content were only weakly correlated with basal area. Δ and P/PE were negatively correlated with leaf nitrogen content. Δ was negatively correlated with leaf density but positively correlated with specific leaf area. This is consistent with the theory that larger leaf nitrogen content and smaller specific leaf area are associated with increased photosynthetic capacity and increased leaf‐scale water‐use‐efficiency, and that Δ is influenced by mesophyll conductance. It is concluded that canopy conductance is a more important determinant of growth in water‐limited conditions than either leaf area index or leaf traits in fertilized stands of E. globulus. Water availability was dictated more by rainfall than soil type.     

不同降水条件下科尔沁沙地南缘疏林草地樟子松针叶δ13C和叶性状特征

通过比较不同自然降水年份(极端干旱和极端湿润)19年生疏林草地樟子松的针叶δ13C、比叶面积和干物质含量,结合土壤含水量和地下水埋深,探讨了极端降水对樟子松水分利用的影响.结果表明:干旱年份(2009)樟子松林土壤含水量显著低于湿润年份(2010),但樟子松当年生针叶的δ13C在两年间没有显著差异,且两年相同月份间亦无显著差异;干旱年份当年生针叶的比叶面积显著低于湿润年份,而不同年份间干物质含量的差异不显著.在两种极端降水条件下,樟子松的水分利用效率没有明显变化,主要通过改变当年生针叶的比叶面积来适应降水量的变化.对于地下水埋深高于3.0m的疏林草地樟子松人工林生态系统,极端干旱不会严重影响樟子松的存活和生长.     

Long-term yield and water use efficiency under various tillage systems in Mediterranean rainfed conditions

Conservation tillage (CT) can be beneficial for soil, water and soil organic matter conservation in Mediterranean areas that are prone to soil erosion and where water availability for crops is the main factor for sustainability. CT is the best option to protect the soil from erosion, improve infiltration, reduce soil evaporation and so conserve rainwater to increase crop water use (WU) and also water use efficiency (WUE). While CT can play an important role in reaching the stability and sustainability of these agricultural systems, performance depends upon the choice and adoption of an appropriate soil management (tillage) system. In rainfed areas of the Ebro Valley, winter cereals are the main crop sown. This paper presents the results of 15 years of research in different soil and climatic conditions of the area of CT on water conservation, WU and WUE. Long‐term experiments, comparing different tillage systems, were established in 1987, 1990 and 1992, at three locations in the Ebro Valley, chosen according to their degree of aridity (Selvanera, Agramunt and El Canós). Results reveal that CT was most effective in increasing yield under the driest conditions at Agramunt (10–15%), still effective with a smaller advantage under slightly wetter conditions at El Canós (5–10%) but ineffective at Selvanera, the wettest site. CT only increased WU in some years at Agramunt and never at the other two sites. The benefits of CT to both increased yield at Agramunt and El Canós were determined by improved WUE arising from changes in the pattern of WU before and after anthesis.     

Reflectance Indices Indicative of Changes in Water and Pigment Contents of Peanut and Wheat Leaves

Measurements of reflectance in visible and near-infrared spectral regions were made on detached leaves of two crop species of different leaf morphology, structure, and water content (peanut and wheat) throughout progressive desiccation. Relative water content (RWC) was well correlated with water index (WI) but even better with the ratio of WI and normalized difference vegetation index. RWC was also significantly correlated with structural independent pigment index indicative of carotenoids/chlorophyll ratio. New indication is thus provided to assess leaf water content and apply simple and fast radiometric techniques for plant water stress management. This revised version was published online in June 2006 with corrections to the Cover Date.     

Provenance effect on carbon assimilation,photochemistry and leaf morphology in Mediterranean Cistus species under chilling stress

• The potential resilience of shrub species to environmental change deserves attention in those areas threatened by climate change, such as the Mediterranean Basin. We asked if leaves produced under different climate conditions through the winter season to spring can highlight the leaf traits involved in determining potential resilience of three Cistus spp. to changing environmental conditions and to what extent intraspecific differences affect such a response.
• We analysed carbon assimilation, maximum quantum efficiency of PSII photochemistry (F_v/F_m) and leaf morphological control of the photosynthetic process in leaves formed through the winter season into spring in C. creticus subsp. eriocephalus (CE), C. salvifolius (CS) and C. monspeliensis (CM) grown from seed of different provenances under common garden conditions.
• Intraspecific differences were found in F_v/F_m for CE and CS. Carbon assimilation‐related parameters were not affected by provenance. Moreover, our analysis highlighted that the functional relationships investigated can follow seasonal changes and revealed patterns originating from species‐specific differences in LMA arising during the favourable period.
• Cistus spp. have great ability to modify the structure and function of their leaves in the mid‐term in order to cope with changing environmental conditions. The F_v/F_m response to chilling reveals that susceptibility to photoinhibition is a trait under selection in Cistus species. Concerning carbon assimilation, differing ability to control stomatal opening was highlighted between species. Moreover, seasonal changes of the functional relationships investigated can have predictable consequences on species leaf turnover strategies.

     

Gene discovery in cereals through quantitative trait loci and expression analysis in water-use efficiency measured by carbon isotope discrimination

Drought continues to be a major constraint on cereal production in many areas, and the frequency of drought is likely to increase in most arid and semi-arid regions under future climate change scenarios. Considerable research and breeding efforts have been devoted to investigating crop responses to drought at various levels and producing drought-resistant genotypes. Plant physiology has provided new insights to yield improvement in drought-prone environments. Crop performance could be improved through increases in water use, water-use efficiency (WUE) and harvest index. Greater WUE can be achieved by coordination between photosynthesis and transpiration. Carbon isotope discrimination (Δ(13) C) has been demonstrated to be a simple but reliable measure of WUE, and negative correlation between them has been used to indirectly estimate WUE under selected environments. New tools, such as quantitative trait loci (QTL) mapping and gene expression profiling, are playing vital roles in dissecting drought resistance-related traits. The combination of gene expression and association mapping could help identify candidate genes underlying the QTL of interest and complement map-based cloning and marker-assisted selection. Eventually, improved cultivars can be produced through genetic engineering. Future efficient and effective breeding progress in cereals under targeted drought environments will come from the integrated knowledge of physiology and genomics.     

Relationships of grain δ13C and δ18O with wheat phenology and yield under water-limited conditions

Stable carbon isotope composition (δ¹³C) of dry matter has been widely investigated as a selection tool in cereal breeding programmes. However, reports on the possibilities of using stable oxygen isotope composition (δ¹⁸O) as a yield predictor are very scarce and only in the absence of water stress. Indeed, it remains to be tested whether changes in phenology and stomatal conductance in response to water stress overrule the use of either δ¹³C or δ¹⁸O when water is limited. To answer this question, a set of 24 genotypes of bread wheat ( Triticum aestivum ) were assayed in two trials with different levels of deficit irrigation and a third trial under rainfed conditions in a Mediterranean climate (northwest Syria). Grain yield (GY) and phenology (duration from planting to anthesis and from anthesis to maturity) were recorded, and the δ¹³C and δ¹⁸O of grains were analysed to assess their suitability as GY predictors. Both δ¹³C and δ¹⁸O showed higher broad-sense heritabilities ( H ²) than GY. Genotype means of GY across trials were negatively correlated with δ¹³C, as previously reported, but not with δ¹⁸O. Both isotopes were correlated with grain filling duration, whereas δ¹⁸O was also strongly affected by crop duration from planting to anthesis. We concluded that δ¹⁸O of grains is not a proper physiological trait to breed for suboptimal water conditions, as its variability is almost entirely determined by crop phenology. In contrast, δ¹³C of grains, despite being also affected by phenology, still provides complementary information associated with GY.     

Variations in life-form composition and foliar carbon isotope discrimination among eight plant communities under different soil moisture conditions in the Xilin River Basin,Inner Mongolia,China

Water is one of the key limiting factors for the survival and growth of plant species in arid and semi-arid steppe regions. Different plant functional groups (PFGs) based on life-forms differ in their strategies to cope with limited water availability. The foliar carbon isotope discrimination () value provides an integrated measurement of internal plant physiological and external environmental properties affecting photosynthetic gas exchange over the time interval when the carbon was fixed. In this study, we surveyed the composition and values of various life-forms (shrubs, sub-shrubs, perennial grasses, perennial forbs and annuals) in eight different plant communities along a soil moisture gradient in the Xilin River Basin, Inner Mongolia, China. Our results showed that: (1) life-forms occurred variously in eight steppe communities with different soil moisture status; (2) in wetter habitats, forbs were more abundant and accounted for the majority of aboveground biomass, whereas grasses became more important in drier habitats. Shrubs and sub-shrubs increased with decreasing soil water availability and their relative biomass rapidly increased in degraded steppe and sand dune communities. (3) The numerical order of the mean values of life-forms is as follows: perennial grasses (15.86) < shrubs (16.10) < perennial forbs (16.45)=annuals (16.41) < sub-shrubs (17.55), reflecting their differences in water use efficiencies. The significant differences in the values among these life-forms suggested that life-form-based PFGs not only represent a morphological classification of these plants, but could also represent a functional group integrating different physiological processes such as water use strategies, which may partially explain the differences in PFG composition and competitive ability of co-existing species along environmental gradients in the Xilin River Basin.     

Genotypic variations in carbon isotope discrimination, transpiration efficiency, and biomass production in rice as affected by soil water conditions and N

Genotypic and environmental (soil water regime and N level) variation in carbon isotope discrimination (CID) in relation to the gas exchange, transpiration efficiency (A/T), and biomass production were investigated in field experiments using eleven rice (Oryza sativa L.) genotypes. The results showed that genotype was more dominant for variation in CID than in total biomass. Genotypic ranking in CID was consistent across environments because of small genotype × environment interactions. Japonica genotypes tended to have lower CID than indica genotypes. Higher soil water and lower N rate significantly increased CID. Variation in CID was slightly smaller for water regime than for genotype. There was a negative correlation between CID andA/T among genotypes within water regimes. Genotypic variation in CID was associated mainly with variation in stomatal conductance under all soil water regimes and with photosynthetic capacity in late growth stages under aerobic soil conditions. The decrease in CID at higher N was probably due to lower stomatal conductance under aerobic soil conditions and to higher photosynthetic rates under submerged soil conditions. The correlation between biomass and CID was not clear in aerobic soil, whereas it was positive in submerged soil, which indicated that the significance of lower or higher CID for improving biomass productivity may differ under different soil water regimes. Overall, the results implied a possible use of CID as a selection criterion for genotypic improvement inA/T and productivity in rice.     

Cytokinin-mediated source/sink modifications improve drought tolerance and increase grain yield in rice under water-stress

Drought is the major environmental factor limiting crop productivity worldwide. We hypothesized that it is possible to enhance drought tolerance by delaying stress-induced senescence through the stress-induced synthesis of cytokinins in crop-plants. We generated transgenic rice (Oryza sativa) plants expressing an isopentenyltransferase (IPT) gene driven by P(SARK) , a stress- and maturation-induced promoter. Plants were tested for drought tolerance at two yield-sensitive developmental stages: pre- and post-anthesis. Under both treatments, the transgenic rice plants exhibited delayed response to stress with significantly higher grain yield (GY) when compared to wild-type plants. Gene expression analysis revealed a significant shift in expression of hormone-associated genes in the transgenic plants. During water-stress (WS), P(SARK)::IPT plants displayed increased expression of brassinosteroid-related genes and repression of jasmonate-related genes. Changes in hormone homeostasis were associated with resource(s) mobilization during stress. The transgenic plants displayed differential expression of genes encoding enzymes associated with hormone synthesis and hormone-regulated pathways. These changes and associated hormonal crosstalk resulted in the modification of source/sink relationships and a stronger sink capacity of the P(SARK)::IPT plants during WS. As a result, the transgenic plants had higher GY with improved quality (nutrients and starch content).     

Trophodynamic linkage between river runoff and coastal fishery yield elucidated by stable isotope data in the Gulf of Lions (NW Mediterranean)

The link between climate-driven river runoff and sole fishery yields observed in the Gulf of Lions (NW Mediterranean) was analysed using carbon- and nitrogen stable isotopes along the flatfish food webs. Off the Rhone River, the main terrestrial (river POM) and marine (seawater POM) sources of carbon differed in ¹³C (–26.11 and –22.36, respectively). Surface sediment and suspended POM in plume water exhibited low ¹³C (–24.38 and –24.70, respectively) that differed more from the seawater POM than from river POM, demonstrating the dominance of terrestrial material in those carbon pools. Benthic invertebrates showed a wide range in ¹⁵N (mean 4.30 to 9.77) and ¹³C (mean –23.81 to –18.47), suggesting different trophic levels, diets and organic sources. Among the macroinvertebrates, the surface (mean ¹³C –23.71) and subsurface (mean ¹³C –23.81) deposit-feeding polychaetes were particularly ¹³C depleted, indicating that their carbon was mainly derived from terrestrial material. In flatfish, ¹⁵N (mean 9.42 to 10.93) and ¹³C (mean –19.95 to –17.69) varied among species, indicating differences in food source and terrestrial POM use. A significant negative correlation was observed between the percentage by weight of polychaetes in the diet and the ¹³C of flatfish white muscle. Solea solea (the main polychaete feeder) had the lowest mean ¹³C, Arnoglossus laterna and Buglossidium luteum (crustacean, mollusc and polychaete feeders) had intermediate values, and Solea impar (mollusc feeder) and Citharus linguatula (crustacean and fish feeder) exhibited the highest ¹³C. Two different benthic food webs were thus identified off the Rhone River, one based on marine planktonic carbon and the other on the terrestrial POM carried by the river. Deposit-feeding polychaetes were responsible for the main transfer of terrestrial POM to upper trophic levels, linking sole population dynamics to river runoff fluctuations.     

No evidence of carbon limitation with tree age and height in Nothofagus pumilio under Mediterranean and temperate climate conditions

Background and Aims

Trees universally decrease their growth with age. Most explanations for this trend so far support the hypothesis that carbon (C) gain becomes limited with age; though very few studies have directly assessed the relative reductions of C gain and C demand with tree age. It has also been suggested that drought enhances the effect of C gain limitation in trees. Here tests were carried out to determine whether C gain limitation is causing the growth decay with tree age, and whether drought accentuates its effect.

Methods

The balance between C gain and C demand across tree age and height ranges was estimated. For this, the concentration of non-structural carbohydrates (NSCs) in stems and roots of trees of different ages and heights was measured in the deciduous temperate species Nothofagus pumilio. An ontogenetic decrease in NSCs indicates support for C limitation. Furthermore, the importance of drought in altering the C balance with ontogeny was assessed by sampling the same species in Mediterranean and humid climate locations in the southern Andes of Chile. Wood density (WD) and stable carbon isotope ratios (δ¹³C) were also determined to examine drought constraints on C gain.

Key Results

At both locations, it was effectively found that tree growth ultimately decreased with tree age and height. It was found, however, that NSC concentrations did not decrease with tree age or height when WD was considered, suggesting that C limitation is not the ultimate mechanism causing the age/height-related declining tree growth. δ¹³C decreased with tree age/height at the Mediterranean site only; drought effect increased with tree age/height, but this pattern was not mirrored by the levels of NSCs.

Conclusions

The results indicate that concentrations of C storage in N. pumilio trees do not decrease with tree age or height, and that reduced C assimilation due to summer drought does not alter this pattern.     

Genotypic and within canopy variation in leaf carbon isotope discrimination and its relation to short-term leaf gas exchange characteristics in cassava grown under rain-fed conditions in the tropics

In a field rain-fed trial with 15 cassava cultivars, leaf gas exchanges and carbon isotope discrimination (Δ) of the same leaves were determined to evaluate genotypic and within-canopy variations in these parameters. From 3 to 7 months after planting leaf gas exchange was measured on attached leaves from upper, middle, and lower canopy layers. All gas exchange parameters varied significantly among cultivars as well as canopy layers. Net photosynthetic rate (P _N) decreased from top canopy to bottom indicating both shade and leaf age effects. The same trend, but in reverse, was found with respect to Δ, with the highest values in low canopy level and the lowest in upper canopy. There were very significant correlations, with moderate and low values, among almost all these parameters, with P _N negatively associated with intercellular CO₂ concentration (C _i), ratio of C _i to ambient CO₂ concentration C _i/C _a, and Δ. Across all measured leaves, Δ correlated negatively with leaf water use efficiency (WUE = photosynthesis/stomatal conductance, g _s) and with g _s, but positively with C _i and C _i/C _a. The later parameters negatively correlated with leaf WUE. Across cultivars, both P _N and correlated positively with storage root yield. These results are in agreement with trends predicted by the carbon isotope discrimination model.     

Prairie plant phenology driven more by temperature than moisture in climate manipulations across a latitudinal gradient in the Pacific Northwest,USA

Plant phenology will likely shift with climate change, but how temperature and/or moisture regimes will control phenological responses is not well understood. This is particularly true in Mediterranean climate ecosystems where the warmest temperatures and greatest moisture availability are seasonally asynchronous. We examined plant phenological responses at both the population and community levels to four climate treatments (control, warming, drought, and warming plus additional precipitation) embedded within three prairies across a 520 km latitudinal Mediterranean climate gradient within the Pacific Northwest, USA. At the population level, we monitored flowering and abundances in spring 2017 of eight range‐restricted focal species planted both within and north of their current ranges. At the community level, we used normalized difference vegetation index (NDVI) measured from fall 2016 to summer 2018 to estimate peak live biomass, senescence, seasonal patterns, and growing season length. We found that warming exerted a stronger control than our moisture manipulations on phenology at both the population and community levels. Warming advanced flowering regardless of whether a species was within or beyond its current range. Importantly, many of our focal species had low abundances, particularly in the south, suggesting that establishment, in addition to phenological shifts, may be a strong constraint on their future viability. At the community level, warming advanced the date of peak biomass regardless of site or year. The date of senescence advanced regardless of year for the southern and central sites but only in 2018 for the northern site. Growing season length contracted due to warming at the southern and central sites (~3 weeks) but was unaffected at the northern site. Our results emphasize that future temperature changes may exert strong influence on the timing of a variety of plant phenological events, especially those events that occur when temperature is most limiting, even in seasonally water‐limited Mediterranean ecosystems.