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Earliness per se (Eps) genes may play a critical role in further improving wheat adaptation and fine-tuning wheat development to cope with climate change. There are only few studies on the detailed effect of Eps on wheat development and fewer on the interaction of Eps with the environment and other genes determining time to anthesis. Furthermore, it seems relevant to study every newly discovered Eps gene and its probable interactions as the mechanisms and detailed effects of each Eps may be quite different. In the present study, we evaluated NILs differing in the recently identified Eps-7D as well as in Ppd-D1 at three temperature regimes (9, 15 and 18 °C) under short day. The effect of Eps-7D on time to anthesis as well as on its component phases varied both qualitatively and quantitatively depending on the allelic status of Ppd-D1 and temperature, being larger in a photoperiod-sensitive background. A more noticeable effect of Eps-7D (when combined with Ppd-D1b) was realised during the late reproductive phase. Consequently, the final leaf number was not clearly altered by Eps-7D, while floret development of the labile florets (florets 2 and 3 in this case, depending on the particular spikelet) was favoured by the action of the Eps-7D-late allele, increasing the likelihood of particular florets to become fertile, and consequently, improving spike fertility when combined with Ppd-D1b. This research was funded by the Spanish Research Agency (AEI), project AGL2015-69595-R, and the International Wheat Yield Partnership (IWYP), project IWYP25FP.

Earliness per se (Eps) genes are reported to be important in fine-tuning flowering time in wheat independently of photoperiod (Ppd) and vernalisation (Vrn). UnlikePpdandVrngenes,Epshave relatively small effects and their physiological effect along with chromosomal position are not well defined. We evaluated eight lines derived from crossing Paragon and Baj (late and early flowering respectively), vernalisation insensitive, to study the detailed effects of two newly identified QTLs,Eps-7DandEps-2Band their interactions under field conditions. The effect of both QTLs were minor but their effect was modulated by the allelic status of the other. While the magnitude of effect of these QTLs on anthesis was similar, they are associated with very different profiles of pre-anthesis development which also depends on their interaction.Eps-7Daffected both duration before and after terminal spikelet while not affecting final leaf number (FLN) soEps-7D-earlyhad a faster rate of leaf appearance.Eps-2Bacted more specifically in the early reproductive phase and slightly altered FLN without affecting the leaf appearance rate. BothEps-7Dand2Baffected the spike fertility by altering the rate of floret development and mortality. The effect of the latter was very small but consistent in that the -lateallele tended to produced more fertile florets.

38 Pags.- 13 Figs.- 2 Tabls. The definitive version is available at: https://www.sciencedirect.com/science/journal/11610301, Genetic gains in wheat yield are required to meet future demand. In order to design strategic crosses, it is essential to quantify the genetic variation among elite cultivars on traits responsible for the adaptation and the generation of yield components. We explored the degree of variation within well-adapted commercial cultivars on developmental phases (vegetative, early reproductive and late reproductive), final leaf number and phyllochron, leaf and spikelet plastochron, number of spikelets, florets, and dynamics of floret primordia development. Variation in flowering time among cultivars was minor but they differed in the partitioning of that time into particular phases. Main differences were found for the length of the late reproductive phase, partly linked to those in final leaf number. Variations on number of fertile florets at anthesis were mostly explained by floret survival, due to differential ability of labile florets to maintain normal development between cultivars. These differences were related to the duration of the floret development, but not to the synchrony of development among florets. These findings provide useful information for breeders when selecting genotypes for strategic crosses, as considering floret survival through duration of floret development in one of the parents (for which these processes shall be characterised in the elite germplasm) might increase the likelihood of transgressive segregation towards high-yielding offspring., Funding was provided by ADAPTAWHEAT (a project funded by the European Commission under the 7th Framework Programme for Research and Technological Development) and by project AGL2012- 35300 funded by the Ministry of Economy and Competitiveness of Spain. HO held a FPI scholarship from the Spanish Ministry of Science and Innovation, and PP held a PhD scholarship from the University of Lleida.

Future grain yield (GY) improvements require the identification of beneficial traits within the context of high yield potential and not just based on the pleiotropic effect of traits such as crop height and heading date. We evaluated 1937 lines from Nested Association Mapping (NAM) population derived from 13 bi-parental varietal crosses under field conditions. We selected 493 lines with similar time to anthesis to that of the two checks used in the study (across and within each family) which reduced the range of plant height in the selected lines. Yield components were measured in these 493 lines from which 231 lines were selected by excluding lines with lowest number of grains so excluded low yielding lines. Later the subset of 231 lines were evaluated in two field experiments (2016−17, CS1 and 2017−18, CS2). Numerical and physiological components of grain yield were measured. The two-step selection maximised GY within an acceptable range of variation for height and anthesis. GY in 231 lines showed very high G × E interaction. Taking both seasons together, we selected lines from upper and lower quartile GY groups to identify stable beneficial trait combinations for improved GY. Differences in GY were explained by grain number driven by increased spike dry weight at anthesis (SDWa) and fruiting efficiency (FE). Increased GY was accompanied by sink limitation. The data points towards increases in grain number as the route towards future GY increases in wheat breeding. Funding for the experimental work was partly provided by projects AGL2015-69595-R, from the Spanish Research Agency (AEI), and IWYP25FP, from the International Wheat Yield Partnership (IWYP). PB held a pre-doctoral research contract from the agency for Management of University and Research (AGAUR) from the Generalitat de Catalunya.

Earlinessper se(Eps) genes are important to fine tune adaptation, and studying their probable pleiotropic effect on wheat yield traits is worthwhile. In addition, it has been shown that some Eps genes interact with temperature. We studied two NILs differing in the newly identifiedEps-7Dbut carrying insensitivePpd-D1 in the background under three temperature regimes (9, 15 and 18 °C) and two photoperiods (12 h and 24 h).Eps-7Daffected time to anthesis as expected and theEps-7D-lateallele extended both the period before and after terminal spikelet. The interaction effect ofEps-7D× temperature was significant but not cross-over: the magnitude and level of significance of the difference between NILs with thelateorearlyallele was affected by the growing temperature (i.e. difference was least at 18 °C and largest at 9 °C), and differences in temperature sensitivity was influenced by photoperiod. Rate of leaf initiation was faster in NIL withEps-7D-earlythan with thelateallele which compensated for the shorter duration of leaf initiation resulting in similar final leaf number between two NILs.Eps-7D-late consistently increased spike fertility through improving floret primordia survival as a consequence of extending the late reproductive phase.

Variations in wheat yield are largely explained by changes in grain number per m2 which is linked to the number of fertile florets at anthesis. This, in turn is the outcome of developmental processes which control floret initiation and mortality. Earliness ‘per se’ (Eps) genes are involved in fine-tuning time to anthesis in wheat (and other cereals) but their effect on development prior to anthesis is less well studied. We aimed to determine effects of Eps genes on spike fertility, quantifying aspects of floret developmental which influence this trait. Field experiments were carried out to record floret primordia generation/degeneration dynamics in near isogenic lines (NILs) with contrasting Eps alleles (late flowering vs early flowering alleles; Eps-late and Eps-early, respectively) derived from the Avalon x Cadenza (AxC) cross with the Eps gene on either chromosome 1D or 3 A and from the Spark x Rialto (SxR) cross with the Eps gene on chromosome 1D. Eps NILs varied in spike fertility: Eps-late alleles increased fertility. Although the effect was in general slight, the magnitude was affected by the particular alleles and the cross used to produce the NILs. Differences in the number of fertile florets were explained by differences in the dynamics of floret development. NILs with Eps-late alleles improved the development of a small number of labile florets allowing them to complete their development to become fertile florets instead of dying, as in lines carrying early alleles. Thus, these alleles improved floret fertility mainly through reducing the rate of floret mortality with no influence on the dynamics of floret primordia initiation or in maximum number of floret primordia. Therefore, Eps genes could be exploited in wheat breeding not only to fine-tune time to anthesis but also to improve spike fertility.

Ppd-1a reduced the number of fertile florets due largely to lowering floret survival, and the effect was not proportional to the duration of the late reproductive phase., As wheat yield is linearly related to grain number, understanding the physiological determinants of the number of fertile florets based on floret development dynamics due to the role of the particular genes is relevant. The effects of photoperiod genes on dynamics of floret development are largely ignored. Field experiments were carried out to (i) characterize the dynamics of floret primordia initiation and degeneration and (ii) to determine which are the most critical traits of such dynamics in establishing genotypic differences in the number of fertile florets at anthesis in near isogenic lines (NILs) carrying photoperiod-insensitive alleles. Results varied in magnitude between the two growing seasons, but in general introgression of Ppd-1a alleles reduced the number of fertile florets. The actual effect was affected not only by the genome and the doses but also by the source of the alleles. Differences in the number of fertile florets were mainly explained by differences in the floret generation/degeneration dynamics, and in most cases associated with floret survival. Manipulating photoperiod insensitivity, unquestionably useful for changing flowering time, may reduce spike fertility but much less than proportionally to the change in duration of development, as the insensitivity alleles did increase the rate of floret development.

Variation in photoperiod sensitivity in wheat plays a major role in the crop adaptation to wide agronomic environments. Photoperiod insensitivity is provided by Ppd-Aa, Ppd-B1a and Ppd-D1a alleles. Effects of the genome, doses and source of the particular Ppd-1a alleles on time to anthesis has not been analysed simultaneously in the same experiments, even less under field conditions; and the effects on particular phases rather than on considering only the total time to anthesis and on phyllochron have not been considered for this range of allele combinations. We carried out field experiments during two consecutive growing seasons to assess differences in time to anthesis, in its component phases, in final leaf number, phyllochron and tillering across wheat isogenic lines differing in specific Ppd alleles and homoeoalleles. In addition to confirming that the introgression of Ppd-1a alleles advanced anthesis time (on average by 307, 251 and 191 °C d if there were 3, 2 or 1 insensitivity alleles introgressed, respectively), we found that the variation between photoperiod insensitive genotypes was largely dependent on the varietal source of Ppd-B1a which could be stronger or weaker than Ppd-D1a depending on the donor considered. All components of time to flowering: the particular sub-phases (P < 0.001) as well final leaf number (P < 0.001) and phyllochron of late-appearing leaves (P < 0.05) were sensitive to Ppd-1a alleles, but the strength of particular alleles on particular components was different, so that similar adjustments in time to anthesis could be achieved with different partitioning of developmental time between the considered phases. We also found that although they did not affect phyllochron of the first 7 leaves, that of the leaves appearing later was consistently reduced in lines carrying Ppd-1a. Tillering was sensitive too, but not final number of spikes due to compensations between tillering and tiller mortality. Funding was provided by ADAPTAWHEAT (a project funded by the European Commission under the 7th Framework Programme for Research and Technological Development) and by project AGL2012-35300 funded by the Ministry of Economy and Competitiveness of Spain. HO held a FPI scholarship from the Spanish Ministry of Science and Innovation and PP held from University of Lleida scholarship. We wish to thank Ignacio Romagosa for valuable advice in the statistical procedures and Andreu Casellas for excellent technical assistance.

In a large comparison of experimental and published data across a wide range of environmental conditions, a cross-over interaction of durum vs . bread wheat yield was found to be the reverse of expectations, where the former would outyield the latter under low-yielding conditions and vice-versa . In this study, we aimed to establish the physiological basis of this species cross-over interaction, analysing whether it was established before or after anthesis and to determine if it was related to differences in sink-strength. Our study indicates that the consistent species-by-environment interaction for yield is the consequence of a few traits related to sink-strength that are determined around anthesis. Thus, differences in the spike weight and N accumulated at anthesis, which produced important variations in the number of grains and presumably generated differences in sink-strength, seemed to be involved. This conclusion is supported by the strong relationship between grain yield and the biomass produced during grain filling, in addition to the fact that the variation in individual grain weight, as well as its rate or the duration of grain filling, was poorly related to yield and the post-anthesis biomass.

Earliness per se (Eps) genes are important to fine tune adaptation, and studying their probable pleiotropic effect on wheat yield traits is worthwhile. In addition, it has been shown that some Eps genes interact with temperature and therefore determining the likely Eps × temperature interaction is needed for each newly identified Eps gene. We studied two NILs differing in the newly identified Eps-7D (carrying insensitive Ppd-D1 in the background) under three temperature regimes (9, 15 and 18 °C) and two photoperiods (12 and 24 h). Eps-7D affected time to anthesis as expected and the Eps-7D-late allele extended both the period before and after terminal spikelet. The interaction effect of Eps-7D × temperature was significant but not cross-over: the magnitude and level of significance of the difference between NILs with the late or early allele was affected by the growing temperature (i.e., difference was least at 18 °C and largest at 9 °C), and the differences caused due to temperature sensitivity were influenced by photoperiod. The rate of leaf initiation was faster in NIL with Eps-7D-early than with the late allele which compensated for the shorter duration of leaf initiation resulting in similar final leaf number between two NILs. Eps-7D-late consistently increased spike fertility through improving floret primordia survival as a consequence of extending the late reproductive phase. This research was funded by Spanish Research Agency (AEI), project AGL2015-69595-R; International Wheat Yield Partnership (IWYP), project IWYP25FP.

In a previous paper (Field Crops Res. 203, 114-127), we showed that the difference in yield potential between a contemporary and a traditional cultivar was due to differences in fruiting efficiency, likely derived from differences in spike fertility (fertile florets per spike) while having similar spike dry weights at anthesis. In this study, we determined the mechanistic bases of these genotypic differences in spike fertility analysing the initiation of all floret primordia per spike (up to 8), the maximum number of florets initiated per spikelet, and the associated floret developmental rates and their fate to become fertile florets under contrasting photoperiod (natural vs extended) and nitrogen availability (50 or 200 kg N ha−1) during the stem elongation phase. Under potential growing conditions (natural photoperiod, high nitrogen availability), the contemporary cultivar owed its higher spike fertility to the improved rate of floret development, which mainly determined an improved level of floret primordia survival to produce fertile florets. The sensitivity of the floret developmental patterns to faster development due to exposure to an extended photoperiod and, to a larger extent, a reduction in N availability was similar for both cultivars, providing a basis for the consistent differences in spike fertility across a range of environments. The response again determined a main effect through increasing floret mortality reducing therefore the level of fertile florets per spikelet in these conditions. Funding was provided by project AGL2009-11964 granted by the Spanish Ministry of Science and Innovation. AF held a FPU scholarship from the Spanish Government.

Efforts to enhance the yield of wheat require increases in the number of grains per m2 (GN) which depends on spike fertility, i.e. the number of fertile florets or grains per spike. Fruiting efficiency (FE), the number of grains set per unit spike dry weight at anthesis, has been proposed as an alternative trait to improve spike fertility and thus GN. The use of FE in realistic breeding programs requires genetic variation within well-adapted, high-yielding modern cultivars and a lack of trade-off with other yield components. We quantified FE variation within Spanish elite material and determined whether the variation implied trade-offs with spike dry weight or grain weight (GW). Field experiments under contrasting nitrogen conditions were carried out with nine elite genotypes (eight modern cultivars and one advanced line) in which the number of fertile florets and grains were determined for different spikelets and the whole spike. Furthermore, the partitioning of dry matter was investigated at anthesis and maturity. Overall, the yield was strongly related to GN because environmental and genetic factors contributed to this relationship, although as expected for elite material the environmental effect was much greater than the genotype. Environmental effects on GN mainly influenced spike dry weight at anthesis (SDWa) whereas most genotypic differences were due to FE variation. Averaging across environments, the FE was c. 110–170 grains gspike−1. Although some G × E interactions were observed, there was no reversal in ranking between genotypes with extreme FEs across environments, and genotypic differences across environmental conditions were consistent (h2 = 0.46). The relationship between FE and the individual weight of fertile florets tended to be negative, although rather than a constitutive reduction in floret size driving the improvements in FE, this relationship appeared to reflect the increased proportion of smaller grains (distal florets in each spikelet) due to changes in spike fertility. FE was not related to grain weight (either the average of all grains or that of the most proximal grains). Therefore, differences in FE among the elite genotypes did not imply constitutively-linked changes in grain weight. When chaff dry weight was used as a proxy for SDWa to estimate genotypic variation in FE, we observed the consistent underestimation of FE and also changes in the ranking of genotypes. Funding was provided partially by ADAPTAWHEAT (a project funded by the European Commission under the 7th Framework Programme for Research and Technological Development) and partially by project AGL2012-35300 funded by the Ministry of Economy and Competitiveness of Spain.

Maize yield has to be increased in the next decades in order to satisfy world demand. This increase has to be achieved in a scenario of climate change particularly characterised by heat stress. Several agronomic and genetic strategies for increased tolerance to high temperatures will be necessary. Some results in other crops showed interactions between high temperature stress and nitrogen (N) availability. To the best of our knowledge, this interaction has not been tested in maize. Therefore, the aim of this study was to explore, under field conditions, whether the magnitude of yield penalty imposed by elevated canopy air temperature around flowering or during early grain filling is affected by N availability. In particular we aimed to (i) quantify the magnitude of yield losses by heat stress around flowering and during early grain filling, (ii) determine whether N fertilisation affects these magnitudes, and (iii) identify whether the effects are indirect (through affecting growth) or directly on the grain set and/or grain growth capacity. Four field experiments were carried out during four consecutive years in NE Spain. The treatments consisted in a factorial combination of one or two hybrids, two or three levels of N fertilisation and three temperature conditions. The temperature treatments consisted of a control (plots grown under natural temperature throughout the growing season) and two treatments in which the temperatures of the canopy were increased in the field for relatively short periods. All experiments were well watered to avoid water stress. The elevated canopy air temperature treatment only increased the maximum temperature in a relatively small magnitude (mean daily temperature was increased at the ears height by c. 1 °C each day of treatment). However, yield penalty imposed by heat stress was in general very noticeable and dramatic when the treatment included the critical period for grain number determination, around silking. The damage was much stronger in the long- than in the short-cycle hybrid. Exposing the plants to elevated canopy air temperature during the critical period reduced harvest index, from values of around 46% in unheated conditions to 20% under elevated canopy air temperature, being the reduction higher under N200 than under N0 fertilisation treatments. As far as we are aware, we showed for the first time in maize grown in field conditions, that the losses in yield in response to elevated canopy air temperature treatment were magnified by the N availability. The effect of N on emphasising the penalties seemed not to be a direct effect of this nutrient but and indirect effect through affecting growth. The effect was through affecting the capacity of the plants to set grains and to a lesser extent to allow grain weight to be maximised; and it was independent of any (potentially additional) effects on either uncoupling anthesis and silking or on pollen amount and viability.

The introgression of awns as a source of photoassimilates during grain filling has received has long been of particular attention in wheat. Although the benefits for average grain weight (AGW) seem consistent, those for yield still remain unclear, and the causes for the improved AGW have not been examined. In this study we carried out three field experiments. In two of them, one in CIMMYT (Mexico) and the other in Lleida (Spain), we compared awned-awnless near isogenic lines (NILs) subjected to defoliation and degraining during the effective period of grain filling, whilst in another experiment we analysed the responses of modern cultivars to awn detachment 10 days after anthesis under two nitrogen levels. There were not consistent differences in yield between the awned-awnless NILs neither across the locations nor across genetic backgrounds. This lack of effect on yield was associated with a compensatory effect of the presence of awns on decreasing grain number while increasing AGW. The manipulation treatments did not consistently affect AGW neither in the NILs nor in the modern cultivars. Focusing on the responses of the NILs, firstly there were some statistically significant responses in AGW but they were both mostly very small and inconsistent between defoliation and degraining treatments as well as between genetic backgrounds. Secondly these responses failed to be consistently more noticeable in the awnless than in the awned lines (as expected if the presence of awns increases AGW through increasing photosynthetic capacity during grain filling). The increase on AGW due to the presence of awns seemed ascribed to both an indirect and a direct effect. The former would simply reflect the consequence of awns reducing grain number: this would naturally be the consequence of an increased failure of distal florets to become fertile which would have reduced the proportion of grains constitutively smaller as well. A direct true effect on the capacity of the grains to grow also emerged with the analysis of individual size of each particular grain for the awned-awnless NILs revealing a constitutively higher potential size in the awned NILs, even for the largest grains. Funding for the experimental work at Lleida was provided by project AGL2015-69595-R from the Spanish Research Agency (AEI). Part of the funding for the experiment was supported by BASF Agricultural Solutions. RSB was supported by Juan de la Cierva program JDC-Formación (FJCI-2016-28164). JWK held a pre-doctoral research contract from the Government of Catalonia. CRA and GM were supported by the Sustainable Modernization of Traditional Agriculture (MasAgro) an initiative from the Secretariat of Agriculture and Rural Development (SADER) and CIMMYT and the International Wheat Yield Partnership (IWYP). JLA acknowledges the support of ICREA Academia, Generalitat de Catalunya, Spain.

High-temperatures reduce yield of wheat and with global warming episodes of heat waves (only few days of high maximum temperatures) during grain filling will become more frequent. It has been recently reported that the magnitude of the yield penalties imposed by high temperatures under field conditions may interact with nitrogen (N) availability both in barley and maize. We determined, under field conditions, the penalties imposed by post-anthesis high-temperatures waves (increased maximum –but not minimum– temperatures during part of the grain filling period) on wheat yield under contrasting soil N supply during two consecutive years. The high temperature treatment was imposed for 10 d starting 10 d after anthesis by placing over the crops transparent polyethylene film (125 μm) mounted on wood structures of 1.5 m height above the ground. This high-temperature and the unheated controls were imposed on 5 modern and well adapted cultivars under contrasting N availabilities (376, 268 and 68 KgN ha−1). Averaged across N conditions, high-temperature treatments reduced yield by c. 1.5 Mg ha−1 (a loss of c. 17%) even though the treatment was rather mild in terms of different average temperature during grain filling. The magnitude of the loss was consistently shaped by the N condition in which the treatment was imposed: yield penalty produced by high-temperature increased from less than 1 to 2.6 Mg ha−1 (which represents losses from 10 to 25%) in parallel with the increased N supply. The penalties were related to both yield components (grain number and average grain weight) which also were more severely penalised under high than under low N supply. As episodes of high-temperature waves will become more frequent in the future the tools used to establish the needs of N fertilisation should be revised as the rates maximising yield (or gross margin) might induce higher sensitivities to these episodes. Also simulation models used to upscale physiological responses to regional or even global domains might need to be revised to include the effect of heat waves (which would be larger per °C increase than what is estimated from experiments increasing temperature during the whole day and over longer periods) as well as the interaction with N supply. Authors thank Semillas Batlle S.A. for their support in the setting up and maintenance of the field plots. Funding was provided by project AGL2012-35300 of the Ministry of Economy and Competitiveness of Spain.

Future increases in grain number may be achieved through improving fruiting efficiency (FE; grains set per unit spike dry weight at anthesis). In recent experiments it was found a negative relationship between the average weight of the grains and FE. The objective of this paper was to determine whether this negative relationship was constitutive (i.e. the increased FE caused all grains to be smaller) or resulted from the increased proportion of grains of smaller potential size. Four experiments, involving 8 different environmental conditions, were carried out during two consecutive growing seasons (2008–2009 and 2009–2010) in which two durum wheat cultivars contrasting in FE and average grain weight were compared. In these conditions we determined FE and carried out a comprehensive study analysing in detail the weights of each individual grain along each of the spikelets. Averaging all experimental conditions, there was a negative relationship between average grain weight (GW) and fruiting efficiency (FE). Donduro with low FE produced in average heavier grains than Vitron. However, when considering individual grains there were, in general, no differences in GW. Therefore, the negative relationship between average grain weight and FE would not be constitutive, and would mostly represent the increase in the proportion of distal grains within the spikelets or of grains from apical spikelets as a consequence of an increased FE, reducing the average size of the grain but mostly unaffecting the size of particular grains. This provides further and stronger support to the idea of using FE as a criterion to further raise yield potential in wheat breeding programmes.

Waterlogging, if occurring within the stem elongation period (SE), is particularly critical for yield determination. We quantified for the first time the effect of waterlogging duration during SE on yield and studied whether the effects were only direct on resource capture or whether there were feed-forward effects as well. We grew wheat (cv. Soissons) outdoors in long tubes (1.25 m deep) forming a normal canopy and imposed different treatments in SE to finish simultaneously around anthesis (0, 4, 8, 12, 16, 20 and 24 days) plus two complementary treatments (8 and 16 days) starting 10 days after the onset of SE. Yield was reduced linearly with the duration of waterlogging c. 2 % dwaterlogging−1. Treatments mainly affected pre-anthesis spike growth reducing the number of fertile florets and grains, not affecting fruiting efficiency. The magnitude of grain number loss was inversely proportional to the hierarchy of the spikes and spikelets. Grain weight was more marginally reduced, likely through the effects on the size of the ovaries of the developing florets. This reveals a direct effect of waterlogging on the capture of resources with no major feed-forward effects. Losses were in agreement with those from other studies for particular durations of waterlogging.

The frequency and intensity of extreme high temperature events are expected to increase with climate change. Higher temperatures near anthesis have a large negative effect on maize (Zea mays, L.) grain yield. While crop growth models are commonly used to assess climate change impacts on maize and other crops, it is only recently that they have accounted for such heat stress effects, despite limited field data availability for model evaluation. There is also increasing awareness but limited testing of the importance of canopy temperature as compared to air temperature for heat stress impact simulations. In this study, four independent irrigated field trials with controlled heating imposed using polyethylene shelters were used to develop and evaluate a heat stress response function in the crop modeling framework SIMPLACE, in which the Lintul5 crop model was combined with a canopy temperature model. A dataset from Argentina with the temperate hybrid Nidera AX 842 MG (RM 119) was used to develop a yield reduction function based on accumulated hourly stress thermal time above a critical temperature of 34 °C. A second dataset from Spain with a FAO 700 cultivar was used to evaluate the model with daily weather inputs in two sets of simulations. The first was used to calibrate SIMPLACE for conditions with no heat stress, and the second was used to evaluate SIMPLACE under conditions of heat stress using the reduction factor obtained with the Argentine dataset. Both sets of simulations were conducted twice; with the heat stress function alternatively driven with air and simulated canopy temperature. Grain yield simulated under heat stress conditions improved when canopy temperature was used instead of air temperature (RMSE equal to 175 and 309 g m−2, respectively). For the irrigated and high radiative conditions, raising the critical threshold temperature for heat stress to 39 °C improved yield simulation using air temperature (RMSE: 221 g m−2) without the need to simulate canopy temperature (RMSE: 175 g m−2). However, this approach of adjusting thresholds is only likely to work in environments where climatic variables and the level of soil water deficit are constant, such as irrigated conditions and are not appropriate for rainfed production conditions. Fil: Gabaldón Leal, C.. Centro Alameda del Obispo; España Fil: Webber, H.. Universitat Bonn; Alemania Fil: Otegui, Maria Elena. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Universidad de Buenos Aires. Facultad de Agronomía. Departamento de Producción Vegetal. Cátedra de Producción Vegetal; Argentina Fil: Slafer, Gustavo Ariel. Universidad de Lleida; España. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Institució Catalana de Recerca i Estudis Avancats; España Fil: Ordóñez, R. A.. Universidad de Lleida; España Fil: Gaiser, T.. Universitat Bonn; Alemania Fil: Lorite, I. J.. Centro Alameda del Obispo; España Fil: Ruiz Ramos, M.. Universidad Politécnica de Madrid; España Fil: Ewert, F.. Universitat Bonn; Alemania

Highlight Outcomes of floret initiation, mortality/survival, grain set/abortion, and fruiting efficiency have been quantified in 30 cultivars and connected to the processes determining the fate of floret primordia using ovary size., Spike fertility traits are critical attributes for grain yield in wheat (Triticum aestivum L.). Here, we examine the genotypic variation in three important traits: maximum number of floret primordia, number of fertile florets, and number of grains. We determine their relationship in determining spike fertility in 30 genotypes grown under two contrasting conditions: field and greenhouse. The maximum number of floret primordia per spikelet (MFS), fertile florets per spikelet (FFS), and number of grains per spikelet (GS) not only exhibited large genotypic variation in both growth conditions and across all spikelet positions studied, but also displayed moderate levels of heritability. FFS was closely associated with floret survival and only weakly related to MFS. We also found that the post-anthesis process of grain set/abortion was important in determining genotypic variation in GS; an increase in GS was mainly associated with improved grain survival. Ovary size at anthesis was associated with both floret survival (pre-anthesis) and grain survival (post-anthesis), and was thus believed to ‘connect’ the two traits. In this work, proximal florets (i.e. the first three florets from the base of a spikelet: F1, F2, and F3) produced fertile florets and set grains in most cases. The ovary size of more distal florets (F4 and beyond) seemed to act as a decisive factor for grain setting and effectively reflected pre-anthesis floret development. In both growth conditions, GS positively correlated with ovary size of florets in the distal position (F4), suggesting that assimilates allocated to distal florets may play a critical role in regulating grain set.

Highlights • Time to terminal spikelet and from then to anthesis were largely independent. • The length of the stem elongation phase was slightly but positively related to grains per m2. • Fruiting efficiency was critical for determining grain number, but it was also negatively related to grain weight. • The length of the stem elongation phase seems to have imposed an upper threshold for fruiting efficiency., Developmental patterns strongly influence spike fertility and grain number, which are primarily determined during the stem elongation period (i.e. time between terminal spikelet phase and anthesis). It has been proposed that the length of the stem elongation phase may, to an extent, affect grain number; thus it would be beneficial to identify genetic variation for the duration of this phase in elite germplasm. Variation in these developmental patterns was studied using 27 elite wheat lines in four experiments across three growing seasons. The results showed that the length of the stem elongation phase was (i) only slightly related to the period from seedling emergence to terminal spikelet, and (ii) more relevant than it for determining time to anthesis. Thus, phenological phases were largely independent and any particular time to anthesis may be reached with different combinations of component phases. Yield components were largely explained by fruiting efficiency of the elite lines used: the relationships were strongly positive and strongly negative with grain number and with grain weight, respectively. Although fruiting efficiency showed a positive trend with the duration of stem elongation that was not significant, a boundary function (which was highly significant) suggests that the length of this phase may impose an upper threshold for fruiting efficiency and grain number, and that maximum values of fruiting efficiency may require a relatively long stem elongation phase.

Yield differences among wheat cultivars and its responsiveness to resource availability are usually related to grain number per m 2 , and further increases in grain number are required, but now beyond differences in time to anthesis and plant height (that are mostly optimised in traditional growing regions). The objective of this paper was to determine and quantify differences in yield physiology between different modern well adapted durum wheats grown in contrasting conditions (in addition we also tested if a higher amount of resources due to removal of competition could affect yield through the same determinants to nitrogen (N) availability). Four experiments were carried out during two growing seasons (2008–09 and 2009–2010); all in all under eight different growing conditions (Gc1–Gc8). In the first season four different modern cultivars were used and in the second season we selected the two highest-yielding cultivars, which differed in their responses to N in terms of spike fertility. Yield was closely related to biomass and N uptake due to both growing conditions and genotypic effects in all experiments. The existence of variation among modern cultivars in these yield determinants is relevant as further increases in yield must be achieved beyond further increases in partitioning. Yield differences among cultivars were based on their differences in grains per m 2 or average grain weight depending on which cultivars were compared. Overall grain number was related to spike dry weight at anthesis, and no evidences were found for a direct involvement of N in grain number determination, beyond the general effect of N availability on spike dry matter. However, this overall effect was mainly driven by growing conditions whilst differences between genotypes seemed associated with either spike dry weight at anthesis or fruiting efficiency, which varied significantly between cultivars. Interestingly, grain weight was consistently higher in the cultivars with lower fruiting efficiency than in cultivars maximising the number of grains set per unit of spike dry weight at anthesis suggesting a constitutive component for the commonly reported negative relationship between grain weight and grain number. In conclusion, the main differences in yield between modern, well adapted wheats were related to their biomass and N uptake and to the sink-strength during post-anthesis, being this strength increased either by having more grains or grains of potentially greater size, depending on the specific cultivar and as a consequence of a trade-off between fruiting efficiency and grain size.

Stem lodging, a permanent displacement of cereal stems which may reduce wheat yields significantly, affects wheat crops in both developed and developing countries, mainly under high-yielding conditions. Although breeding has decreased these losses by reducing the likelihood of the plant lodging through reduced height (mainly by the introgression of dwarfing genes), lodging continues to affect wheat yields. We aimed to quantify how wheat breeding has modified the sensitivity of yield to lodging, identifying physiological causes for yield reductions. Two field experiments were carried out in Mediterranean Spain during the 2006/07 and 2007/08 growing seasons. The first season two genotypes (Aragon 03 and ID-2151) and two lodging treatments (control and lodged near anthesis) were analysed. The second season four genotypes (Aragon 03, Estrella, Anza and ID-2151) and three lodging treatments (control, lodged from either jointing to maturity or anthesis to maturity) were imposed. In addition, in the 2007/08 experiment the source–sink ratios of five representative spikes of each plot were modified through hand trimming all spikelets from one side of each spike soon after anthesis. Yield losses due to early lodging were reduced by wheat breeding: yield penalties in the oldest and newest genotypes were 61% and 43% (mean values of both lodging treatments), respectively while it was 57% in intermediate cultivars (mean value of both lodging treatments and cultivars). Lodging decreased both the number of grains per m 2 and the average grain weight (AGW). The decrease of the number of grains per m 2 was associated with reductions in the crop growth, while the AGW loss seemed related to both a reduction of the availability of assimilates to fill grains and a direct effect on grain size potential. Lodged plants exhibited stronger source-limitation during grain filling than those un-lodged, while the response of the grain weight to sink-strength reductions in lodged plants was very similar to that in the un-trimmed controls, indicating that grain filling in un-lodged and in strongly lodged canopies was similarly balanced in terms of source–sink ratios.

Further increasing yield potential remains one of the main objectives of wheat breeding, even in stressful environments. In general, past genetic gains were associated with increases in harvest index, and future gains should be related to greater biomass. Identifying genetic sources for such improvement may be relevant. Researchers of TRITIMED identified DH lines of durum wheat apparently possessing not only high yield potential but also good yield stability. We aimed to determine physiological attributes responsible for yield and stability among a set of genotypes derived from two parents (Cham 1 and Lahn) and four of the most promising lines of the DH population (2401, 2408, 2410, 2517). Seven field trials were carried out within the Mediterranean agricultural region of the Ebro Valley, under a wide range of conditions (ca 2–10 mg ha−1). In four of these experiments, sub-plots were included with source-sink manipulations imposed after anthesis. Cham 1, a cultivar known for high yields in semi-arid conditions, showed the highest yield potential. Although it showed less yield stability than Lahn, even under the lowest yielding conditions its yield was not significantly lower than that of Lahn. RILs 2408, 2410, 2004 and 2517 slightly outyielded Lahn in high-yielding conditions, but under poorer environments they tended to yield less. Interestingly, yield differences were closely related to their biomass rather than harvest index. Thus yield differences relating to the number of grains per m2 were due to differences in spike dry matter at anthesis, reflecting in part genotypic differences in crop growth from jointing to anthesis. In general grain weight did not respond to spike trimming after anthesis, although in two experiments the grain weight of Cham 1 did so. Thus, even the highest-yielding cultivar possessed grains that overall seemed more limited by its constitutive capacity to grow than by the availability of resources to reach this capacity (though occasionally they may be co-limited). Overall, the most interesting feature was the empirical evidence that improvement of biomass within elite material is a worthwhile objective.

The stem elongation phase seems critical in yield potential determination in barley ( Hordeum vulgare L.). Extending its duration, without modifying total time to anthesis, has been proposed as a promising breeding tool. A prerequisite for its use is that the duration of phases before and after jointing (that including leaf and spikelet initiation, LS, and that of stem elongation, SE) should be under different genetic control. In addition, little is known about the implications of changes in the duration of LS and SE upon other developmental traits which could affect other aspects of yield generation, such as phyllochron and tillering. Thus, the objectives of the present work were to study the genetic variability in LS and SE, in traits related to leaf appearance and tillering, as well as their relationships, in a double-haploid (DH) population derived from the cross Henni × Meltan. DH lines and both parents were studied in four field trials. Despite the similarity in development between parents, there was significant genetic variability in duration of both LS and SE ( i.e . considerable transgressive segregation was observed), with no major genetic correlations found between them. Although some significant genetic correlations were found between duration of phases and leaf appearance and tillering traits, it has been shown that modifying the duration of LS does not necessarily imply concomitant changes in traits that could be important for an early expansion of the crop canopy (i.e. phyllochron, onset and rate of tillering).

Past breeding achievements in grain yield were mainly related to increases in harvest index (HI) without major changes in biomass production. As modern cultivars have already high HI, future breeding to improve grain yield will necessarily focus on increased biomass. Improved biomass would depend on our capacity to improve the amount of photosynthetically active radiation intercepted by the crop (IPAR%) or the efficiency with which the canopy converts that radiation into new biomass (radiation use efficiency, RUE). Four field experiments with a set of wheat cultivars selected, bred and introduced in the Mediterranean area of Spain and that represent important steps in wheat breeding in Spain were conducted in order to identify whether and how wheat breeding in this area affected the amount of IPAR% and RUE both before and after anthesis. Although there was genotypic variability, cultivars did not show any consistent trend with the year of release of the cultivars for their biomass, pre and post-anthesis IPAR%, Crop growth rate (CGR) or RUE but, the post-anthesis CGR and RUE of the two oldest genotypes were lower than that of the other cultivars. As the oldest genotypes have lower number of grains per m2 than their modern counterparts, it is suggested that post-anthesis RUE in these cultivars was reduced by lack of sinks and therefore further increases in grains per unit area in modern cultivars could permit to improve biomass via increases in post-anthesis RUE.

The critical period for grain yield determination has not been determined for triticale. We aimed to identify it, determining the relative importance of both the major yield components and the dry matter acquisition by the spikes at anthesis. A field experiment was carried out with two triticales, differing in tillering capacity, subjected to shading treatments at five different timings from early tillering to maturity. Results showed that reductions in grain yield were more significant when shading was imposed during 3 weeks before and 1 week after anthesis. Reductions in grain yield by shading treatments were associated with lower number of grains per m2 more than with changes in the average grain weight. Reductions in grains per m2 were due to reductions in the number of fertile florets per spike, affecting grains per spike. The assimilate acquisition by the spikes during the critical period was a key determinant of floret survival. Grain number per m2 was related with photothermal quotient during 30 days before anthesis and spike dry weight at anthesis, though the goodness of the prediction compared with wheat, was lowered by poorer grain setting percentage.

Three field experiments comparing bread wheats representing different eras of genetic improvement in the Mediterranean area of Spain (one landrace, seven cultivars released since 1950 and two advanced breeding lines) were carried out at the province of Lleida (Catalonia, north-eastern Spain). Experiments were conducted during the 2004/05, 2005/06 and 2006/07 growing seasons at Gimenells and during 2005/06 at Foradada in order to quantify gains in yield, and in its physiological determinants, during much of the last century. At Gimenells, the first two seasons were conducted under relatively high-yielding backgrounds (Gimenells, irrigated) and in the last season under moderately stressed conditions (Gimenells, rainfed), while at Foradada the experiment was conducted under a severely stressed low-yielding environment. For almost all the traits analyzed in this paper, there were only significant differences when cultivars were grown under relatively high-yielding environments. In this conditions, grain yield increased with the year of release of the cultivars from 1940 to the 1970s, with no clear further improvements thereafter. On the other hand, total biomass did not show any association with the year of release of the cultivars. Results showed that, even when breeding was performed under Mediterranean conditions, harvest index was the main attribute responsible for yield improvements. Stem height was linearly and negatively related with harvest index, implying a marked height reduction until the 1970s, with no clear further reductions thereafter. In general yield was lineally and positively associated with number of grains per m2, while average grain weight did not exhibit any clear trend with the year of release of the cultivars. The increase in number of grains was more associated with that in grains per spike than with differences in spikes per m2. Finally, the increase in number of grains per m2 was associated with both number of grains per unit of spike dry weight at anthesis, or “fruiting efficiency”, and spike dry weight at anthesis.

A better understanding of responsiveness of time to anthesis to water stress in triticale and wheat might help to explain their differences in performance under stress and to identify which attributes or components of time to anthesis are most affected. Three experiments were carried out during the 2004 and 2005 growing seasons to (i) evaluate genotypic variability in phenology responsiveness to drought in triticale and to (ii) explore the variation in the duration of pre-anthesis phases occurring before and after the onset of stem elongation as well as their responsiveness to water stress. Important variation was found among triticale cultivars on time to anthesis, allowing classification of cultivars into long, intermediate and short cycle. Differences in the duration of the stem elongation phase (i.e. from terminal spikelet initiation (TS) to anthesis (ANT)) were as high as 200 °Cd for cultivars with similar time to flowering. Although water stress reduced the time to anthesis (in average by ca. 125 °Cd), cultivars showed a differential responsiveness to drought in phases occurring before or after TS. Some cultivars that were highly sensitive to drought in the emergence (Eme)–TS phase were insensitive in the phase TS–ANT and vice-versa. Water shortage tended to reduce the grain filling period and was most evident when a Mediterranean environment treatment was simulated.

Spike growth during the stem elongation phase is critical for the establishment of the number of fertile florets (FF). A longer period of spike growth would lead to an increased number of FF and grains in wheat. The aim of this study was: (i) to analyze in three different commercial cultivars the impact of a photoperiod extension (NP + 10) during the stem elongation phase on the number of FF and (ii) to determine whether the photoperiod extension effect might be reversed by increasing the availability of assimilates by a direct modification of the source/sink ratio at booting. The number of FF per spike was reduced by extended photoperiod in the experiment sown in an optimum date, though no differences were observed when sown much later, probably due to the poor environmental conditions of a spring sowing. The detrimental effect of lengthening the photoperiod on the number of FF per spike was reversed by trimming the spike at booting in two of the three cultivars analyzed. This reversion, together with the effect of photoperiod extension on FF associated with similar effects on spike growth, may be considered as a confirmation that, at least in part, the detrimental effects of photoperiod extension on FF would be mediated by changes in assimilate supply to the developing florets. No differences were observed between photoperiod treatments on the onset of rapid spike growth; however, extended photoperiod reduced the duration of spike growth phase without differences in the maximum rate of spike growth in two of the three cultivars. Although a positive association was found between the number of FF and spike dry weight at anthesis, differences among cultivars were observed in this relationship, suggesting different carbohydrates partitioning (i.e., FF per unit of spike weight) among them.

An early prediction of crop biomass at maturity and yield is important in different circumstances. The use of spectral reflectance indices, such as the Normalized Difference Vegetation Index (NDVI), has been proposed as a fast, nondestructive way of estimating crop growth capacity. In this study, we examined whether NDVI assessment relatively early in the crop cycle may be useful for predicting final biomass and yield in wheat. To that end, NDVI was measured and biomass quantified regularly from tillering to maturity for six different wheat genotypes grown under a contrasting range of N and water availabilities. In addition, final biomass and yield were measured at maturity. In line with expectations from the literature, we found that NDVI at milk-grain stage was well correlated to final yield and biomass. However, it was also observed that NDVI at the onset of stem elongation was also reasonably correlated to both attributes. Because crop growth in wheat from the end of tillering to anthesis is related to the determination of grain number and yield, we propose the use of NDVI at the onset of stem elongation as a complementary criterion for establishing the required late crop management (N fertilisation, irrigation) practices.

The stem elongation phase in wheat [Triticum aestivum (L.)] is considered critical for yield determination. A longer duration of this phase could hypothetically increase grain set and therefore yield. Genetic variation in the relative duration of the stem elongation phase having been reported, the aim was to pinpoint whether this variability was associated with sensitivity to photoperiod, vernalizing temperatures or to differences in intrinsic earliness. Pairs of cultivars identified as having different duration of the stem elongation phase (from the appearance of the first visible node to anthesis) were grown under natural (short) or extended photoperiod, with or without vernalization. Variability in the duration of this phase, in the cultivars analysed, was related to different sensitivity to photoperiod, while differences in the previous phases were related to sensitivity to both photoperiod (though different to the sensitivity of the following phase) and vernalization.

The stem elongation phase in wheat (Triticum aestivum, L.) and barley (Hordeum vulgare) is important for yield determination as it is the phase when stems and spikes grow actively. A longer duration of this phase could hypothetically increase assimilate availability, fertile florets at anthesis and final grain number. Argentine commercial wheat and barley cultivars were studied to analyse the range of genetic variability existing in the duration of the stem elongation phase, in particular, considering cultivars with similar time to anthesis. It was found that cultivars differing in time to anthesis, not only differed in the length of their early, mostly vegetative phases but also in the stem elongation phase (range 301-535 °C day for wheat and 252-404 °C day for barley), and that the partitioning of a similar time to anthesis into phases was substantially different between cultivars. Various studies with other cultivars were re-analysed, showing that the stem elongation phase did present genetic variability as well. As a preliminary step to identifying the genetic causes behind this variability, an approximation to the possible photoperiod and/or vernalization sensitivities of this large group of Argentine cultivars was presented, by comparing behaviour in two contrasting sowing dates. This has proved useful to breeders since there have been no other previous studies including this large number of commercial cultivars.

In temperate cereals are commonly accepted that determination of grain number (GN) and grain weight (GW) scarcely overlap during the crop cycle. However, the assumption that GW is determined exclusively after anthesis needs to be critically reviewed in the light of reports published over the few years where temperature treatments imposed before anthesis decreased GW of bread wheat. Although these evidences suggest that both GW and GN could be affected by environmental conditions before anthesis little is known about the effect of pre-anthesis temperature on these two main yield components in wheat, barley and triticale at field conditions. In addition, the effect of temperature on GW and GN at different stages prior to anthesis has been scarcely evaluated. The objectives of the current study were: (i) to evaluate the overall response, and specific differences, of GN and GW to pre-anthesis temperature, and (ii) to study the effect of different timings of high temperature at pre-anthesis on GN and GW in wheat, barley and triticale. Three fully irrigated field experiments were carried out in three successive seasons. At each season, a wheat, barley and triticale high yielding cultivar was evaluated at three temperature regimes: control, and two timings of heating before anthesis. During the first and second seasons, the timings of heating were booting-anthesis and heading-anthesis. In the thirst season, the timings were beginning of stem elongation-booting and booting-anthesis. Plots were arranged in a split-plot design with three replicates, where the main plot was assigned to thermal regime and the sub-plots to crop species. To apply heat, transparent chambers equipped with thermostatically controlled electric heaters were used. The thermal regime was controlled by sensors connected to a temperature regulator and recorded using data loggers. Temperature within the chambers was stable across developmental stages, crops, and seasons; it averaged 5.5 °C higher than air temperature. Thermal treatments consistently reduced grain yield (p < 0.05), the magnitude of the effect ranged between 5 and 52%. The highest effect was found when temperature increased during stem elongation (yield decrease: 46%), lowest when treatments were imposed during heading-anthesis (15%) and intermediate for treatments imposed during booting-anthesis (27%). Most effects of thermal treatments on yield were due to parallel effects on GN. However, thermal treatments significantly (p < 0.05) decreased GW during the three seasons. The most effecting treatment on GW was when the crops were heated during the B-A period, i.e. GW decreased up to 23%.

Average grain weight is a major yield component contributing to its variation, especially in Mediterranean regions where grain weight is frequently exposed to terminal stresses affecting grain growth. Most of the literature agrees that wheat grain growth is hardly limited by the source. However, no source–sink ratios studies seem to have been conducted in the Mediterranean region to determine to what degree wheat grain growth is actually limited by the source in these particular regions. We conducted two field experiments in Catalonia (north-eastern Spain), where an old cultivar (Anza) and a more recently released one (Soissons) were sown in a range of different nitrogen and water availabilities and sowing dates. This was to analyse the degree of source limitation for grain growth. Sink size was modified by removing half of the spikelets c . 10 days after anthesis, virtually doubling the availability of assimilates per grain effectively growing. Trimming the spikes did not produce significant changes in grain growth rate or duration of grain filling. Consequently, grain weight did not respond noticeably to the reduction in sink demand and any eventual response has been far from representing a strong competition among grains during grain filling.

Lengthening the late reproductive phase (LRP) of stem elongation in wheat (Triticum aestivumL.), by changing its photoperiod sensitivity independently of the preceding phases, would improve the yield potential through increasing spike weight and the number of fertile florets at anthesis. This paper presents results of a two-year field experiment designed to determine the impact of Ppd-D1and Ppd-B1on (i) the duration of three pre-anthesis developmental phases, and (ii) spike weight and the number of fertile florets at anthesis under two photoperiods during the LRP (natural and an extension of six hours over that). Near isogenic lines of Mercia and single chromosome recombinant lines of Cappelle Desprez were used. Under natural photoperiod, Ppd-D1hastened time to anthesis ca. 500∘C d in both backgrounds by reducing each of the three pre-anthesis phases. Ppd-B1hastened the time to anthesis under natural photoperiod by 178∘C d, mainly by reducing the early reproductive phase. The response to photoperiod of the LRP under extended daylength depended on the Ppdlocus present: Ppd-D1was insensitive while Ppd-B1and the recessive controls were sensitive. For all lines, photoperiod treatments and years, the number of fertile florets was associated with spike dry weight at anthesis (R 2≅ 80%, p< 0.01) which, in turn, was positively related to the intercepted radiation accumulated during the LRP (R 2 45%, p< 0.05). Changing the duration of the LRP through extended photoperiod or through Ppd-D1produced similar results in both backgrounds and years. Thus, altering the duration of the LRP by manipulating photoperiod sensitivity may be an alternative to changing the fertile floret number in wheat. Nevertheless, as no particular allele was responsible for the photoperiod sensitivity only during the LRP, new alleles should be studied to identify the control of photoperiod sensitivity of individual phases to fine-tune the pre-anthesis wheat development.

Grain protein content is one of the most important attributes in bread making quality. Several studies have reported that the type of, and the balances between, storage protein (i.e. gliadins and glutenins) are also relevant. Generally, the modern cultivars present less grain protein content than their predecessors, suggesting that breeding may have reduced baking quality while improving yield. Furthermore, there is little information on whether, and eventually how, breeding influences the types of protein synthesized in the grains, and the sensitivity of the type of protein and baking quality to changes in the environment. This study aimed to determine the stability in baking quality in wheat cultivars released at different eras to variation in the duration of the pre-anthesis period without changes in the sowing date. The experiment studied the combination of three cultivars released in 1940, 1964 and 1994 at two different durations from sowing to anthesis (because of exposure to different photoperiods during stem elongation in the field). Modern cultivars showed lower grain protein content than the oldest ones (11 vs. 14 %). There were no correlation between grain protein content and baking quality (assessed by Zeleny test), indicating that protein type is more important than grain protein content. Gliadins were more variable than glutenins because of differences between cultivars and flowering dates. Wheat breeding also seemed to have improved the stability of quality variables in response to environmental variation during the pre-anthesis period (when yield potential is being determined).

Summary The final yield of a crop is the product of growth during the growing season and a number of developmental processes occurring throughout the life cycle of a crop, with most genes influencing the final outcome to a degree. However, recent advances in molecular biology have developed the potential to identify and map many genes or QTLs related to various important traits, including yield, plant adaptation and tolerance to stresses. Significant G×E interactions for yield have been identified, as have interactions associated with QTLs for yield. However, there is little evidence available to confirm that a QTL for yield from a parental line in one mapping population may improve yield when transferred into an adapted, high-yielding line of another population. In order to narrow the apparent gap between the genotype and the phenotype with regard to yield, it is important to identify key traits related to yield and then attempt to identify and locate the genes controlling them. The partitioning of the developmental time to anthesis into different phases: from sowing to the onset of stem elongation and from then to anthesis, as a relatively simple physiological attribute putatively related to yield, is discussed. If the relationship holds in a wider range of conditions and the genetic factors responsible are located then the genetic basis of yield should be identified. There has also been significant progress in crop simulation modelling. Using knowledge of crop physiology and empirical relationships these models can simulate the performance of crops, including the G×E interactions. Such models require information regarding the genetic basis of yield, which are included in the form of genetic coefficients. Essentially models are constructed as decision-making tools for management but may be of use in detecting prospective traits for selection within a breeding programme. Problems associated with this approach are discussed. This review discusses the need to use crop physiology approaches to analyse components of yield in order to reliably identify the genetic basis of yield.

The manipulation of photoperiod sensitivity during stem elongation (terminal spikelet initiation—anthesis) has been suggested as a means to increase wheat yield potential. Nevertheless, photoperiod sensitivity and the interaction with vernalization during that phase have not been previously explored under field conditions. Thus, a field study was carried out (i) to assess whether duration of stem elongation is sensitive to photoperiod experienced only during that phase, and if so (ii) to evaluate whether vernalization interacts with this photoperiod response and (iii) to determine if altering duration of stem elongation may influence wheat yield and its components. The experiment was a factorial combination of two vernalization treatments (15 (V15) or 50 (V50) days at 4±1 °C and 8 h photoperiod), two photoperiod regimes during stem elongation (natural daylength in the field (NP+0) and one daylength extension of 6 h (NP+6)) and three cultivars (Buck Manantial (BM), Eureka Ferrocarril Sur (EFS), and ProINTA Puntal (PP)). The stem elongation phase responded to the actual photoperiod experienced during that phase in the three cultivars, increasing its duration with shorter photoperiod. In the vernalization-sensitive cultivars EFS and PP, plants vernalized 50 days showed greater response to photoperiod than those vernalized 15 days. In the most photoperiod-sensitive cultivar (BM), as duration of stem elongation lengthened by exposure to less inductive photoperiod, a higher number of fertile florets at anthesis was achieved leading to an increased grain number and thereby to a higher wheat yield. These results reinforce the idea that a longer duration of stem elongation may offer an alternative avenue to improve yield potential in wheat.

A longer duration of stem elongation (from terminal spikelet initiation to anthesis) would result in a higher wheat yield potential as a consequence of the increase in both spike dry weight and number of fertile florets at anthesis. It is not clearly understood, however, which processes are involved in the increased spike dry weight and whether this variable accounts for the higher number of fertile florets when duration of stem elongation is modified by photoperiod treatments. As vernalization interacts with photoperiod during the stem elongation phase, a better understanding of the role of vernalization is also required. A field experiment was carried out using two wheat cultivars (Buck Manantial (BM) and Eureka Ferrocarril Sur (EFS)) subjected to diverse periods of vernalization (V 15 and V 50 , plants vernalized for 15 and 50 days, respectively) and exposed to different photoperiods (NP+0 and NP+6, natural and 6 h extended photoperiod, respectively) only during the stem elongation phase. Floret development together with stem and spike growth during the stem elongation phase were studied. When the stem elongation phase was exposed to short photoperiod (i) duration of spike growth increased without any change in its growth rate, and (ii) most of the spike growth occurred during minimum stem growth rate as the onset of maximum stem growth rate was delayed. In EFS, the vernalization-sensitive cultivar, the former (i) and (ii) was true only when vernalization requirements were met. For both cultivars, the survival of the initiated floret primordia under short photoperiod increased, independently of vernalization. Both partitioning of assimilates to the spike and a direct photoperiod effect seemed to have been involved in the survival of floret primordia when the stem elongation phase was exposed to short photoperiod.

Photoperiod sensitivity is an important feature of flowering time regulation, which enables wheat plants to adapt to a wide range of environments. Although some work has been done on how time to heading or flowering respond to photoperiod in relation to particular Ppd alleles, there is little evidence on whether these alleles contribute to responses at different phases and to associated yield component generation. The aims of this paper were: (i) to analyse the effects of photoperiod on substitution lines with contrasting Ppd alleles, in terms of duration of particular phases, (ii) to determine if there is any relationship between these alleles and the parameters of photoperiod response (photoperiod sensitivity, optimum photoperiod and basic length of the phase), and (iii) to analyse the effects of different photoperiods applied before and after the onset of terminal spikelet on yield component generation. The effects of length and timing of photoperiod extensions on these traits were analysed under field conditions in Chinese Spring and two substitution lines differing in photoperiod sensitivity. Although time to anthesis was similar in the three genotypes in photoperiods longer than 14.5 h, they did differ in their response to photoperiod in particular phases. Sensitivity to photoperiod for daylengths shorter than 14.5 h was also markedly different. The number of leaves generated was affected by photoperiod, determining the duration of the phase from emergence to floral initiation (EM-FI). The length of the phase from floral initiation to terminal spikelet (FI-TS) was determined by the number of spikelets generated and their rate of initiation, which was also affected by photoperiod. The terminal spikelet to anthesis phase (TS-ANT) was only affected by photoperiod in the most sensitive genotype, in which direct photoperiod effects, other than the effects on leaf number and phyllochron, were evident. There was no apparent relation between photoperiod response parameters such as basic length of the phase (Lb) and optimum photoperiod (Po) and particular Ppd alleles.

Although individual grain weight is an important source of variation forgrain yield, there is still poor understanding of the causes determining finalgrain weight. Almost all studies conducted for understanding thedeterminants of grain weight have been focused on the post-anthesis period.However, there is important evidence that pre-anthesis conditions couldalso modify final grain weight. Three experiments including different sowingdates, genotypes and temperature regimes between booting and anthesis,were carried out in Argentina and Mexico to analyse the effect oftemperature and associated traits during the pre- and post-anthesis periodson grain weight under field conditions. In these experiments final grainweight could not be explained by average or maximum temperature duringthe post-anthesis period. However, average temperature between bootingand anthesis was closely related to the observed grain weight differences,probably as a consequence of the effects of this factor on carpel growth.Differences in grain weight between genotypes and grain position weresuccessfully explained by differences in carpel weight at anthesis. Theseresults suggest that our knowledge to determine grain weight could improveif the immediately pre-anthesis period conditions were taken into account.

It has been hypothesised that wheat yields may be increased by lengthening the duration of the stemelongation phase. This paper reports studies on the effects of chromosomes carrying major photoperiod genes (Ppd-A1, Ppd-B1, Ppd-D1) in different genetic backgrounds, on responses to photoperiod before and after jointing, when the onset of stem elongation occurs, and on number of grains per spike. A field experiment considered the effects of two photoperiods on Chinese Spring and 12 substitution lines, in which chromosomes 2A, 2B or 2D had been substituted by those from four contrasting cultivars. The phase from seedling emergence to jointing (EM-JO) was more responsive than that from jointing to anthesis (JO-ANT), but no relationship was found between the duration of these phases. EM-JO length affected leaf and spikelet number and consequently grains per spike, but this component was further influenced by JO-ANT duration. Our results confirmed that the phases are independent in sensitivity, supporting the hypothesis that genetic manipulation of phase duration could enhance yield, but no evidence was found of any particular Ppd allele being responsible for major responses to photoperiod during stem elongation.

The effects of Rht1 and Rht2 alleles on the dynamics of floret development in isogenic lines (dwarf, DD; semi-dwarf, SD and standard height, SH) of spring wheat were investigated. Studies were conducted on wheat grown in the field in each of 4 years and where water and nutrients were non-limiting. The number of grains per spike was significantly greater in the lines with Rht alleles than in the SH lines. Grain number for each line was such that DD>SD>SH. Grains per spike varied with the number of grains per spikelet rather than number of spikelets per spike. Grains per spikelet in turn varied with the number of fertile florets at anthesis. Florets were considered fertile when male and female reproductive organs had developed green anthers and bifidum stigma, respectively. The dwarfing genes had no effect on the percentage of fertile florets setting grain. Increased number of fertile florets per spikelet due to the presence of Rht1 and Rht2 alleles in the genome was a consequence of the higher number of relatively distal primordia, to progress to the stage of fertile floret at anthesis in the DD and SD than in the SH lines. This ability to allow that a greater proportion of distal florets maintain a normal rate of development was related to the fact that Rht alleles produced a more favourable assimilate partitioning to the spike during the pre-anthesis period associated with the reduction in stem growth imposed by Rht alleles. This allowed a higher proportion of the later-initiated floret primordia to produce fertile florets at anthesis.

Field experiments were conducted during 1991 and 1992 to identify the effect of wheat breeding on crop biomass production and its physiological determinants, i.e. radiation interception and radiation-use efficiency. To address this objective, biomass accumulation and its physiological attributes of seven cultivars (including a commercial hybrid) released between 1920 and 1990 were compared in fertilised and irrigated plots. Weeds, pests, diseases and lodging were controlled or prevented. Crop radiation interception was calculated from measurements of incident and transmitted radiation at different times after sowing. Above-ground dry matter was determined at particular stages of development of the cultivars. Radiation-use efficiencies and crop growth rates were calculated for each cultivar for pre-anthesis (in both years) and post-anthesis (in 1992) periods. Biomass at anthesis tended to be less in most recently released cultivars than in older materials. Accumulated intercepted radiation at similar developmental stages also differed between cultivars. These differences, as well as the trend of biomass at anthesis, were caused by differences in the length of developmental phases between cultivars rather than by changes in the architecture of the canopies. The most recently released cultivars had shorter seedling emergence-anthesis periods than older cultivars. Moreover, all cultivars had similar canopy light extinction coefficients ( k ), pre-anthesis radiation-use efficiencies, and crop growth rates. After anthesis, the two oldest cultivars accumulated the least biomass and their radiation-use efficiencies and crop growth rates were smaller than those of modern cultivars. Remarkably, modern cultivars maintained during post-anthesis almost the same levels of crop growth rates and radiation use efficiencies reached during the pre-anthesis period.

Although it has been generally recognized that the difference in yield potential amongst wheat cultivars released in different eras is related to differences in their reproductive sink strength, there have been few investigations about changes in source–sink ratios as a consequence of wheat breeding. In the present study, two field experiments, in which plots were fertilized and irrigated and lodging and diseases were prevented, were carried out with seven cultivars (including a commercial hybrid) representing different periods of plant breeding in Argentina from 1920 to 1990. The cultivars were defoliated during post-anthesis to analyse the response of grain weight at particular positions within the spike (which have intrinsic differences in potential size).Individual grain weight was virtually unaffected by defoliation in the old cultivars, but modern cultivars exhibited a significant reduction in individual grain weight for several positions within the spike, although this reduction was small (c. 15%) and many grains were unaffected. In addition, no relationship was found between individual grain weight in the controls and its reduction due to defoliation.We concluded that if the source–sink ratio is further reduced, the grain yield of modern wheats will be simultaneously limited by the source and the sink. Future breeding should therefore attempt to improve simultaneously both sink and source strengths.

The effects of Rht alleles on root growth and distribution in isogenic lines of spring wheat (Triticum aestivum L.) are described under different environmental conditions. Above-ground biomass, root length, root dry-weight and their distribution along the soil profile were measured by destructive sampling for growth of aerial biomass and extraction of soil cores containing roots. Field experiments were conducted under non-limiting water and nutritional conditions during two consecutive years, using an early and a late sowing date each year. Dwarfing genes significantly reduced plant height and above-ground biomass at anthesis. In addition, stem mass ratio also was reduced with increases in the allelic dosage. Conversely, total root length and root dry-weight per unit area at anthesis were increased with decreased plant height, therefore, root mass ratio tended to be negatively correlated with plant height. Differences in distribution of root length and root dry-weight through the soil profile among lines were largely confined to the upper soil layers (i.e. the top 30 cm). Differences in root dry-weight were more important than in root length, so that the dwarf line had the highest root mass per unit root length. Furthermore, a significant positive correlation between the root mass ratio and stem mass per unit stem length was found. It is suggested that increases in root mass per unit root length associated with Rht alleles are evidencing a surplus of photoassimilates during stem elongation which are used for thickening the roots due to the lack of alternative sinks. Agronomic implications of this effect are discussed.

Three near isogenic lines of spring wheat grown to determine whether Rht dwarfing alleles alter radiation interception. A field study (involving two sowing dates in two growing seasons) with lines containing different allelic dosage of Rht1 and Rht2 (i.e. dwarf, DD; semi-dwarf, SD and tall SH), was conducted without water and nutritional deficiencies. Dwarfing genes did not modify the timing of occurrence of phenological events. Above-ground biomass at anthesis was reduced by 22% in the DD line in relation of the rest of the lines. However, at maturity accumulated biomass of the DD and SH lines were not significant different. Dwarfing genes increased the light attenuation coefficient (k, with values of 0.48, 0.62 and 0.78 for the SH, SD and the DD line respectively). A similar trend was followed to the leaf thickness (estimated by the specific leaf weight, SLW). Despite the differences observed among the lines, both in k and SLW values, they did not differ significantly in the proportion of incoming radiation intercepted by the canopy, nor in the cumulative intercepted radiation during the pre and post-anthesis periods. Radiation use efficiency (RUE) differed significantly among the lines. While RUE during pre-anthesis was the lowest in the DD line, from anthesis to maturity the lines with Rht alleles showed higher RUE values than the SH line. The low pre-anthesis RUE in the DD lines could be associated with (i) poor canopy architecture due to reductions in leaf sheath and internode lengths and/or (ii) reduced canopy CO2 exchange rate. Post-anthesis RUE was lower than that recorded pre-anthesis in all lines. But the magnitude of the reduction was inversely related to the doses of the Rht alleles. Post-anthesis RUE appeared to be closely and positively associated with the number of grains set per unit biomass at anthesis. This relationship suggests a regulatory effect of the sink size on the efficiency of the crop to convert radiation into biomass during this period.

A better understanding of the effects of different source?sink ratio during the grain filling period on grain growth may be relevant in order to further increase cereal grain yield. The main objective of the present work was to determine the effect of different manipulations of the source?sink ratios in wheat and barley grown at four different environmental conditions on responsiveness of sinks (grain growth and yield) and sources (spike photosynthesis and water soluble carbohydrates in the stems). Four treatments were imposed 7 days after anthesis in two contrasting locations with low- and high-inputs conditions in wheat (cv. Soissons) and barley (cv. Sunrise): they were a control, a treatment removing all the spikelets from the upper half of the spikes (TS), and shadings decreasing incident radiation by 75% on the whole canopy (SW) or only on the leaves (having the top area of the meshes individual holes for each spike to be exposed to solar radiation, SL). As expected grain yield was closely related to grain number per m2. Average grain weight was reduced by shading treatments far more markedly in Sw than in SL. Interestingly, significant amounts of water soluble carbohydrates in the stems remained at maturity in SL and Sw treatments and spike photosynthesis in SL was consistently higher than in the unshaded controls in both species. These results may be an indication that wheat and barley are not source-limited during grain filling and that only when subjected to an extremely severe stress, grain size would be reduced due to lack of enough assimilates available to fill them. Fil: Serrago, Roman Augusto. Universidad de Buenos Aires. Facultad de Agronomía. Departamento de Producción Vegetal. Cátedra de Cerealicultura; Argentina Fil: Alzueta, Ignacio. Universidad de Buenos Aires. Facultad de Agronomía. Departamento de Producción Vegetal. Cátedra de Cerealicultura; Argentina Fil: Savin, Roxana. Universitat de Lleida. Departament de Producció Vegetal i Ciència Forestal; España. Universitat de Lleida. Agrotecnio; España Fil: Slafer, Gustavo Ariel. Universitat de Lleida. Departament de Producció Vegetal i Ciència Forestal; España. Institució Catalana de Recerca i Estudis Avancats; España. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura; Argentina

Wheat yield depends on the number of grains per square metre, which in turn is related to the number of fertile florets at anthesis. The dynamics of floret generation/degeneration were studied in contrasting conditions of nitrogen (N) and water availability of modern, well-adapted, durum wheats in order to understand further the bases for grain number determination. Experiments were carried out during the 2008-2009 and 2009-2010 growing seasons at Lleida (NE Spain). The first experiment involved four cultivars (Claudio, Donduro, Simeto, and Vitron) and two contrasting N availabilities (50 kgN ha(-1) and 250 kgN ha(-1); N50 and N250) while experiment 2 included the two cultivars most contrasting in grain setting responsiveness to N in experiment 1, and two levels of N (N50 and N250), under irrigated (IR) and rainfed (RF) conditions. In addition, a detillering treatment was imposed on both cultivars under the IR+N250 condition. The number of fertile florets at anthesis was increased by ~30% in response to N fertilization (averaging across treatments and spikelet positions). The effect of N and water availability was evident on floret developmental rates from the third floret primordium onwards, as these florets in the central spikelets of all genotypes reached the stage of a fertile floret in N250 while in N50 they did not. In this study, clear differences were found between the cultivars in their responsiveness to N by producing more fertile florets at anthesis (through accelerating developmental rates of floret primordia), by increasing the likelihood of particular grains to be set, or by both traits. We thank the team of the crop physiology laboratory of the UdL for technical assistance. Funding was provided by the Spanish Ministry of Science and Innovation through project AGL2009- 11964. AF held a FPU, AP2006-03719, scholarship from the Spanish Ministry of Education.

A wheat cultivar (Condor) was grown in two experiments (thermal regimes 18/13 and 21/16°C) under low (298 μE m-2: s-1) radiation regimes during either an early phase from seedling emergence to terminal spikelet initiation (S2), a late phase from terminal spikelet initiation to anthesis (S2), or for the full period from seedling emergence to anthesis (S12), or high (560 μE m-2s-1) radiation throughout the growing period (S12) to determine whether developmental events are affected by radiation. The main developmental events considered in this study were the timing of terminal spikelet initiation and anthesis, the final number of leaf and spikelet primordia initiated in the apex and the rate of leaf appearance. Number of grains per spike and culm height were also measured. The duration of each phenophase was not affected by radiation intensity. Temperature affected the rate of wheat development, but the acceleration of development due to temperature during the seedling emergence - terminal spikelet initiation phase only slightly reduced (from 24.8 to 23.2 days). Differences in time from terminal spikelet initiation to anthesis were greater than in the earlier phases, having been the duration reduced from 24.6 to 20.0 days due to high temperature. Associated with the lack of effect of radiation on phasic development and the negligible effect of temperature on the duration of the early phases of development, final Leaf number was practically unchanged in this study by either the radiation level or the growing temperature. Thus, radiation did not affect the rate of leaf initiation. The number of spikelets was affected by neither the treatments nor the thermal environment. The rates of leaf appearance were accelerated by temperature. Radiation, on the other hand, did not significantly alter the rates of leaf appearance in any of the treatments. As expected from many sources in the literature, the number of grains per spike was significantly affected by radiation during the phase from terminal spikelet initiation to anthesis. Due to the lack of significant effects of radiation on the developmental patterns of wheat, the changes in number of grains per spike were due to changes in the number of grains born in each spikelet. The results of the present study were compared with others available in the literature on the effects (or lack of them) of radiation and CO2 concentration on phasic development, plastochron and phyllochron in wheat to reach the general conclusion that the rate of developmental events in wheat, in contrast to other plants, is almost completely independent of the availability of assimilates, with a possible exception for the Equatorial latitudes.