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.