Sink limitations to yield in wheat : how could it be reduced?

Further genetic gains in wheat yield are required to match expected increases in demand. This may require the identification of physiological attributes able to produce such improvement, as well as the genetic bases controlling those traits in order to facilitate their manipulation. In the present paper, a theoretical framework of source and sink limitation to wheat yield is presented and the fine-tuning of crop development as an alternative for increasing yield potential is discussed. Following a top-down approach, most crop physiologists have agreed that the main attribute explaining past genetic gains in yield was harvest index (HI). By virtue of previous success, no further gains may be expected in HI and an alternative must be found. Using a bottom-up approach, the present paper firstly provides evidence on the generalized sink-limited condition of grain growth, determining that for further increases in yield potential, sink strength during grain filling has to be increased. The focus should be on further increasing grain number per m2, through fine-tuning pre-anthesis developmental patterns. The phase of rapid spike growth period (RSGP) is critical for grain number determination and increasing spike growth during pre-anthesis would result in an increased number of grains. This might be achieved by lengthening the duration of the phase (though without altering flowering time), as there is genotypic variation in the proportion of pre-anthesis time elapsed either before or after the onset of the stem elongation phase. Photoperiod sensitivity during RSGP could be then used as a genetic tool to further increase grain number, since slower development results in smoother floret development and more floret primordia achieve the fertile floret stage, able to produce a grain. Far less progress has been achieved on the genetic control of this attribute. None of the well-known major Ppd alleles seems to be consistently responsible for RSGP sensitivity. Alternatives for identifying the genetic factors responsible for this sensitivity (e.g. quantitative trait locus (QTL) identification in mapping populations) are being considered.

Cultural control of herbicide-resistant Lolium rigidum Gaud. populations in winter cereal in Northeastern Spain

Lolium rigidum Gaud. is one of the most common weed species in winter cereals in Northeastern Spain. Herbicide resistance has been growing since the mid 90's and exclusive herbicide use is not enough in many cases, so that it is necessary to combine as many control tools as possible. Six field trials have been conducted during the cropping seasons 2001-02, 2002-03 and 2003-04 on winter cereal infested with herbicide resistant L. rigidum in Northeastern Spain testing different cultural control strategies. Sowing delay was conducted at five fields, mouldboard ploughing at four fields, the combination of sowing delay and ploughing at two fields, increasing the cereal sowing density and combined with sowing delay at one field. Sowing delay was confirmed to have an irregular efficacy depending on the L. rigidum emergence during the delay period. In the trials, weed emergence was reduced up to 88% in the best case but there was no effect in two cases. Ploughing had a more constant efficacy and reduced weed emergence between 50 and 80% although stoniness impeded in one occasion a correct soil inversion causing a very low efficacy. Increasing the cereal sowing rate did not reduce the weed population. The combination of the different methods did not increase significantly the individual efficacy, and one method was clearly more effective than the other, depending on the trial. In fields with high L. rigidum density, these methods are not effective enough and need to be combined with other methods, which are discussed in the text.

Use of nitrification inhibitor DMPP to improve nitrogen recovery in irrigated wheat on a calcareous soil

The 3,4-dimethyilpyirazole phosphate (DMPP), commercialized as Entec, is a nitrification inhibitor developed by BASF (Germany) that may help to minimize N losses and to obtain a higher profit from N fertilizers. A two-year field trial was established in 2001 in the Northeast of Spain to assess the effects of DMPP on N use efficiency (NUE) and to determine the economic returns. Seven treatments have been carried out comparing the effect of DMPP on pig slurry and on mineral fertilizers. The application of DMPP resulted in better efficiency indexes on mineral fertilizers. An apparent nitrogen recovery of 0.465 kg kg-1, on average, was obtained for the Entec treatment. A net benefit of € 809 ha-1, on average, was obtained for the Entec treatment compared with € 607 ha-1 for the control treatment. The results of this study suggest that the nitrification inhibitor could improve farmer profit in irrigated wheat on a calcareous soil.

Nitrogen replacement value of alfalfa to corn and wheat under irrigated Mediterranean conditions

In crop rotations that include alfalfa (Medicago sativa L.), agronomic and environmental concerns mean that it is important to determine the N fertilizer contribution of this legume for subsequent crops in order to help to increase the sustainability of cropping systems. To determine the N fertilizer replacement value (FRV) of a 2-yr alfalfa crop on subsequent crops of corn (Zea mays L.) followed by wheat (Triticum aestivum L.) under irrigated Mediterranean conditions, two 4-yr rotations (alfalfa-corn-wheat and corn-corn-corn-wheat) were conducted from 2001 to 2004 in a Typic Xerofluvent soil. Corn yields were compared after two years of alfalfa and a third year of corn under monoculture and wheat yields were also compared after both rotations. Corn production after alfalfa outyielded monoculture corn at all four rates of N fertilizer application analyzed (0, 100, 200 and 300 kg N/ha). The FRV of 2-yr alfalfa for corn was about 160 kg N/ha. Wheat grown after the alfalfa-corn rotation outyielded that grown after corn under monoculture at both the rates of N studied (0 and 100 kg N/ha). The FRV of alfalfa for wheat following alfalfa-corn was about 76 kg N/ha. Soil NO3 -N content after alfalfa was greater than with the corn monoculture at all rates of N fertilizer application and this higher value persisted during the second crop after alfalfa. This was probably one of the reasons for the better yields associated with the alfalfa rotation. These results make a valuable contribution to irrigated agriculture under mediterranean conditions, show reasons for interest in rotating alfalfa with corn, and explain how it is possible to make savings when applying N fertilizer.

Survival of wild adults of Ceratitis capitata (Wiedemann) under natural winter conditions in north east Spain

The overwintering of the Mediterranean fruit fly (medfly), Ceratitis capitata (Wiedemann) at the northern limits of its geographic distribution is not yet well known. With the aim of estimating the survival rate of medfly adults in northeast Spain under natural winter conditions, a two-winter-season trial was carried out. A control was carried out in a climatic chamber at 25°C. The results showed that medfly adults were unable to survive the entire winter season in the Girona area. Climatic conditions, including the daily minimum temperature, daily maximum temperature and the high rainfall, appeared to be involved in adult mortality in winter.

Does ear C sink strength contributes to the overcoming of photosynthetic acclimation of wheat plants exposed to elevated CO2?

Wheat plants (Triticum durum Desf., cv. Regallo) were grown in the field to study the effects of contrasting [CO2] conditions (700 versus 370 μmol mol−1) on growth, photosynthetic performance, and C management during the post-anthesis period. The aim was to test whether a restricted capacity of sink organs to utilize photosynthates drives a loss of photosynthetic capacity in elevated CO2. The ambient 13C/12C isotopic composition (δ13C) of air CO2 was changed from-10.2 in ambient [CO2] to-23.6 under elevated [CO2] between the 7th and the 14th days after anthesis in order to study C assimilation and partitioning between leaves and ears. Elevated [CO2] had no significant effect on biomass production and grain filling, and caused an accumulation of C compounds in leaves. This was accompanied by up-regulation of phosphoglycerate mutase and ATP synthase protein content, together with down-regulation of adenosine diphosphate glucose pyrophosphatase protein. Growth in elevated [CO2] negatively affected Rubisco and Rubisco activase protein content and induced photosynthetic down-regulation. CO2 enrichment caused a specific decrease in Rubisco content, together with decreases in the amino acid and total N content of leaves. The C labelling revealed that in flag leaves, part of the C fixed during grain filling was stored as starch and structural C compounds whereas the rest of the labelled C (mainly in the form of soluble sugars) was completely respired 48 h after the end of labelling. Although labelled C was not detected in the δ13C of ear total organic matter and respired CO2, soluble sugar δ13C revealed that a small amount of labelled C reached the ear. The 12CO2 labelling suggests that during the beginning of post-anthesis the ear did not contribute towards overcoming flag leaf carbohydrate accumulation, and this had a consequent effect on protein expression and photosynthetic acclimation.

Evidence for the influence of the North Atlantic Oscillation on the total ozone column at northern low latitudes and midlatitudes during winter and summer seasons.

The strong influence of the winter North Atlantic Oscillation (NAO) on the total ozone column (TOC) in the Northern Hemisphere has been reported in a number of previous studies. In this study we show that this influence is not restricted to the winter season but is also significant in summer. Especially interesting effects of the summer NAO (SNAO) on the TOC are observed over the eastern Mediterranean region, where a strongly positive SNAO index is related to the creation of a geopotential height-negative anomaly over Greece with maximum amplitude at 200 hPa. Another anomaly was observed west of the Iberian Peninsula with similar effects on the TOC. Analyzing 26 years of Total Ozone Mapping Spectrometer (TOMS) and Ozone Monitoring Instrument (OMI) data from the equator to midlatitudes (60°) in the Northern Hemisphere, we demonstrate that the SNAO accounts for up to 30% of the TOC variability with a strong latitudinal and longitudinal dependence. Additionally, we obtain significant correlations between the NAO index and the thermal tropopause pressure and also with the geopotential heights at 200 and 500 hPa. Finally, some indirect connections between NAO and the TOC through teleconnections are also discussed.

The effects of depleted, current and elevated growth [CO2] in wheat are modulated by water availability

Drought is the main constraint on wheat yield in Mediterranean conditions. The photosynthesis, chlorophyll fluorescence and plant growth parameters of durum wheat (Triticum turgidum, L. var. durum) were compared at three [CO2] (i.e., depleted 260 ppm, current 400ppm and elevated 700 ppm) in plants subjected to twowater regimes (i.e.,well-wateredWW, and mildwater stress by drought orwater deficit WS), during pre-anthesis, post-anthesis and the end of grain filling. We showed that [CO2] effects on plants are modulated by water availability. Plants at depleted [CO2] showed photosynthetic acclimation (i.e., up-regulation) and reduced plant biomass and Harvest Index, but depleted [CO2] combined with WS has a more negative impact on plants with decreases in C assimilation and biomass. Plants at elevated [CO2] had decreased plant growth and photosynthesis in response to a down-regulation mechanism resulting from a decrease in Rubisco and N content, but plants exposed to a combination of elevated [CO2] and WS were the most negatively affected (e.g., on plant biomass).

Foliar photochemical processes and carbon metabolism under favourable and adverse winter conditions in a Mediterranean mixed forest, Catalonia (Spain)

Evergreen trees in the Mediterranean region must cope with a wide range of environmental stresses from summer drought to winter cold. The mildness of Mediterranean winters can periodically lead to favourable environmental conditions above the threshold for a positive carbon balance, benefitting evergreen woody species more than deciduous ones. The comparatively lower solar energy input in winter decreases the foliar light saturation point. This leads to a higher susceptibility to photoinhibitory stress especially when chilly (< 12 C) or freezing temperatures (< 0 C) coincide with clear skies and relatively high solar irradiances. Nonetheless, the advantage of evergreen species that are able to photosynthesize all year round where a significant fraction can be attributed to winter months, compensates for the lower carbon uptake during spring and summer in comparison to deciduous species. We investigated the ecophysiological behaviour of three co-occurring mature evergreen tree species (Quercus ilex L., Pinus halepensis Mill., and Arbutus unedo L.). Therefore, we collected twigs from the field during a period of mild winter conditions and after a sudden cold period. After both periods, the state of the photosynthetic machinery was tested in the laboratory by estimating the foliar photosynthetic potential with CO2 response curves in parallel with chlorophyll fluorescence measurements. The studied evergreen tree species benefited strongly from mild winter conditions by exhibiting extraordinarily high photosynthetic potentials. A sudden period of frost, however, negatively affected the photosynthetic apparatus, leading to significant decreases in key physiological parameters such as the maximum carboxylation velocity (Vc,max), the maximum photosynthetic electron transport rate (Jmax), and the optimal fluorometric quantum yield of photosystem II (Fv/Fm). The responses of Vc,max and Jmax were highly species specific, with Q. ilex exhibiting the highest and P. halepensis the lowest reductions. In contrast, the optimal fluorometric quantum yield of photosystem II (Fv/Fm) was significantly lower in A. unedo after the cold period. The leaf position played an important role in Q. ilex showing a stronger winter effect on sunlit leaves in comparison to shaded leaves. Our results generally agreed with the previous classifications of photoinhibition-tolerant (P. halepensis) and photoinhibitionavoiding (Q. ilex) species on the basis of their susceptibility to dynamic photoinhibition, whereas A. unedo was the least tolerant to photoinhibition, which was chronic in this species. Q. ilex and P. halepensis seem to follow contrasting photoprotective strategies. However, they seemed equally successful under the prevailing conditions exhibiting an adaptive advantage over A. unedo. These results show that our understanding of the dynamics of interspecific competition in Mediterranean ecosystems requires consideration of the physiological behaviour during winter which may have important implications for long-term carbon budgets and growth trends.

Comparative performance of the stable isotope signatures of carbon, nitrogen and oxygen in assessing early vigour and grain yield in durum wheat

The present paper studied the performance of the stable isotope signatures of carbon (δ13C), nitrogen (δ15N) and oxygen (δ18O) in plants when used to assess early vigour and grain yield (GY) in durum wheat growing under mild and moderate Mediterranean stress conditions. A collection of 114 recombinant inbred lines was grown under rainfed (RF) and supplementary irrigation (IR) conditions. Broad sense heritabilities (H2) for GY and harvest index (HI) were higher under RF conditions than under IR. Broad sense heritabilities for δ13C were always above 0·60, regardless of the plant part studied, with similar values for IR and RF trials. Some of the largest genetic correlations with GY were those shown by the δ13C content of the flag leaf blade and mature grains. Under both water treatments, mature grains showed the highest negative correlations between δ13C and GY across genotypes. Flag leaf δ13C was negatively correlated with GY only under RF conditions. The δ13C in seedlings was negatively correlated, under IR conditions only, with GY but also with early vigour. The sources of variation in early vigour were studied by stepwise analysis using the stable isotope signatures measured in seedlings. The δ13C was able to explain almost 0·20 of this variation under RF, but up to 0·30 under IR. In addition, nitrogen concentration in seedlings accounted for another 0·05 of variation, increasing the amount explained to 0·35. The sources of variation in GY were also studied through stable isotope signatures and biomass of different plant parts: δ13C was always the first parameter to appear in the models for both water conditions, explaining c. 0·20 of the variation. The second parameter (δ15N or N concentration of grain, or biomass at maturity) depended on the water conditions and the plant tissue being analysed. Oxygen isotope composition (δ18O) was only able to explain a small amount of the variation in GY. In this regard, despite the known and previously described value of δ13C as a tool in breeding, δ15N is confirmed as an additional tool in the present study. Oxygen isotope composition does not seem to offer any potential, at least under the conditions of the present study.