The role of ear photosynthesis in grain filling was studied in a number of durum wheat (Triticum turgidum var durum L.) landraces and varieties from the Middle East, North Africa, and from the collections of ‘Institut National de la Recherche Agronomique’ (INRA, France) and ‘Centro International de Mejora de Maiz y Trigo’ (CIMMYT, Mexico). Plants were grown in the field in a Mediterranean climate. Flag leaves (blade plus sheath) and ears were kept in the dark from 1 week after anthesis to maturity which reduced grain weight by 22.4% and 59.0%, respectively. In a further experiment, the carbon isotope discrimination ratio (Δ) of ear bracts, awns and flag leaves was measured on samples taken at anthesis and on mature kernels. The mean value of Δ for the water soluble fraction of bracts (17.0‰) and awns (17.7‰) were lower than those of leaves (19.5‰) and fairly similar to those of kernels (17.4‰) averaged across all genotypes. Data indicate that most of the photosynthates in the grain come from ear parts and not from flag leaves. In addition, a higher water use efficiency (WUE) of ear parts than of the flag leaf is suggested by their lower Δ values. Gas exchange in ears and flag leaves was measured during grain filling. Averaged over all genotypes, CO2 diffusive conductance was about five times higher in the flag leaf than in the spike (with distal portions of awns outside the photosynthetic chamber) 2 weeks after anthesis. In absolute terms, the dark respiration rate (Rd) was greater than the net photosynthesis rate (Pn) by a factor of 1.74 in the spike, whereas Rd was much smaller, only 22.1, 65.7 and 24.8% of Pn in blade, sheath and awns, respectively. Data indicate that photosynthesis, and hence the water use efficiency (photosynthesis/transpiration), is greatly underestimated in ears because of the high rates of respiration which diminish the measured rates of net CO2 exchange. Results of 13C discrimination and gas exchange show that genotypes from North Africa have higher WUE than those from the Middle East. The high Rd values of ears as well as their low diffusive conductance suggest that CO2 from respiration may be used as source of carbon for ear photosynthesis. In the same way, the anatomy of glumes, for example, supports the role of bracts using internal CO2 as source of photosynthesis. In the first experiment, the Δ in mature grains from culms with darkened ears compared with control culms provided further evidence in support of this hypothesis. Thus, the Δ from kernels of control plants was 0.40 higher than that from ear‐darkened plants, probably because of some degree of refixation (recycling) of respired CO2 in the grains.

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Carbon isotope discrimination (∆) was determined for kernels of six-row barley and durum wheat cultivated in the western Mediterranean basin during the last seven millennia. Samples came from different archaeological sites in Catalonia (north-east Spain) and in the south-east of Spain (mainly eastern Andalusia). Samples from the present were also analysed. Mean values of ∆ for barley and durum wheat grains decreased slightly from Neolithic (7000-5000 BP) to Chalcolithic-Bronze (5000-3000 BP) and Iron ages (3000-2200 BP) both in Catalonia and in south-east (SE) Spain. Values were consistently lower in SE Spain than in Catalonia throughout these five millennia, which suggests that Catalonia was less arid than SE Spain in this period. Within a given region, current discrimination values for kernels of the same cereal species cultivated under rainfed conditions were lower than those of archaeological grains, which implies more arid conditions at present. Furthermore, an empirical relationship between ∆ of mature kernels and total precipitation (plus irrigation where applicable) during grain filling (r 2 ϭ 0.73, N ϭ 25) was established for barley, currently cultivated at different locations in the western Mediterranean basin in Spain. The resulting relationship was applied to the ∆ data for barley kernels from 10 archaeological sites in Catalonia and 10 sites in SE Spain, to estimate the precipitation during grain filling at the time the kernels were produced. For both regions, current climatic conditions are consistently more arid than those inferred for the Neolithic, Bronze and Iron ages. In addition, although Catalonia was estimated to have had consistently wetter conditions (about 20% more precipitation) than SE Spain throughout these millennia, differences in precipitation between these two regions have recently increased, with 79% more precipitation in Catalonia. Results indicate a more rapid increase in aridity in SE Spain than in Catalonia, probably produced during the last few centuries, and due to anthropogenic causes.Abbreviations ∆, stable carbon isotope discrimination; δ 13 C, ratio of CO 2 ; WUE, water-use efficiency; BP, before present; NE Spain, north-east of Spain; SE Spain, south-east of Spain; 13 C/ 12 C relative to PeeDee belemnite standard; p i /p a, ratio of intercellular to atmospheric partial pressure of VPD, vapour pressure deficit.

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The analysis of carbon isotope discrimination (∆) in crop plant remains from archaeological sites may help to assess water availability for early agriculture. This study presents the analysis of ∆ 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 ∆ values greater than 16 ‰, reaching 20 ‰ for some samples of flax. For wheat, ∆ 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 ∆ of mature kernels and total precipitation (plus irrigation where applicable) from heading to maturity (r 2 ⍧ 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 ∆ 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 ∆ 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.

Summary -Two near-isogenic 6-rowed barley genotypes (Hordeum vulgare L cv Atlas), with and without awns, were grown in pots under Mediterranean conditions. The plants were watered daily. For each genotype, total net photosynthesis and water-use efficiency (WUE: net photosynthesis/transpiration) of ears and flag leaves were measured 3 times during grain filling: 1, 3 and 4 weeks after anthesis. The total photosynthesis of awned ears was markedly higher than that of awnless ears, until 3 weeks after anthesis. Thereafter, photosynthetic rates decreased faster in awned than in awnless ears. The WUE of awned ears was always higher than that of awnless ears. In fact, the WUE of awns alone was 2 to 3 times higher than the WUE of awnless ears during grain filling.

Abstract. Epidermal (non‐stomatally‐controlled) conductance from the fourth leaf, first node leaf, flag leaf and ear of durum wheat (Triticum turgidum var durum L.) grown under Mediterranean field conditions has been measured, along with leaf stomatal frequency and the amount and distribution of epicuticular waxes. Measurements were carried out on varieties and land‐races from the Middle East, North Africa, ‘Institut National de la Recherche Agricole’ (INRA) and ‘Centra Internacional de Mejora de Maiz y Trigo’ (CIMMYT). Significant differences were observed among genotypes in the epidermal conductances (ge) of the four organs. For each of the four organs tested, genotypes from the Middle East and CIMMYT showed higher ge. values than those from North Africa and INRA. Ears showed epidermal conductances that were more than four times higher than those of leaves when ge. values were expressed per unit dry weight. The amount of epicuticular waxes was higher in the fourth leaves, intermediate in the first node and flag leaves and lower in the ears. For each organ, ge differences among genotypes were unrelated with the amount of epicuticular waxes. Removal of epicuticular waxes by dipping the organs into chloroform significantly increased the epidermal conductance for the fourth and first node leaves and the ear. However, this did not occur for the flag leaf. For the fourth leaf, ge of intact leaves and ge of leaves in which epicuticular waxes were removed were unrelated (r = ‐0.265). The regression coefficient of this relation for the first node and flag leaves showed values of 0.666 and 0.650 (P > 0.05), respectively, and values were even higher in the ear (r > m 0.892, P > 0.01). Scanning electron microscope analysis showed that wax bloom decreased from the fourth leaf to the flag leaf, whereas the extent of amorphous wax increased. Wax bloom in leaves consisted mainly of deposits of thin wax plates. In the ears and the adaxial surface of flag leaves, fibrillar waxes predominated. In the first node and flag leaves, the wax deposits on the adaxial side cover the surface of the leaf more densely and uniformly than those on the abaxial side. There was no significant correlation between ge and total stomatal density, or between ge and either adaxial or abaxial stomatal density for any sample of the three different leaves. The contribution of epicuticular waxes plus total stomatal frequency only explained 42.4, 11.8, 28.3 and 16% of ge (per unit leaf area) variations for the fourth leaf, first node leaf, flag leaf and the combined variation of the three leaves together, respectively. From these results, it is concluded that complex interrelationship between different morphophysiological characteristics probably control ge differences among genotypes and that these interrelationships differ for each different plant part.

The mercury (Hg) accumulation mechanism was studied in rape (Brassica napus) plants grown under a Hg concentration gradient (0 mM-1,000 mM). Hg mainly accumulated in roots. Therefore, the presence of phytochelatins (PCs) was studied in the roots of the plants. The high stability of the PC-Hg multicomplexes (mPC-nHg) seems to be the main reason for the lack of previous Hg-PC characterization studies. We propose a modification of the method to detect and quantify unbound PC of Hg in plant extracts via high-performance liquid chromatography coupled to electrospray tandem mass spectrometry and inductively coupled plasma mass spectrometry in parallel. We separated the PC from the Hg by adding the chelating agent sodium 2,3-dimercaptopropanesulfonate monohydrate. We only detected the presence of PC after the addition of the chelating agent. Some multicomplexes mPC-nHg could be formed but, due to their large sizes, could not be detected. In this study, only PC 2 was observed in plant samples. Hg accumulation was correlated with PC 2 concentration (r 2 5 0.98).