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Delayed canopy senescence or ‘stay-green’ (SG) trait in maize (Zea mays L.) could improve drought tolerance. Two field trials comparing four to six maize hybrids with different senescence rate were carried out at Buenos Aires, Argentina, varying water availability during the reproductive period. Green leaf area at maturity was related to kernel weight (r2 = 0.94***) but its relationship with yield was weaker (r2 = 0.51–53*) and post-silking dry matter remobilisation was negatively related with the SG trait (r2 = 0.84**). Two additional experiments were carried out in 63 L pots by withholding irrigation in half of the pots after silking. The SG hybrid achieved lower root biomass at silking, a shallower root distribution and larger root growth in the post-silking period. Under drought conditions, stomatal conductance was lower in the SG hybrid but photosynthetic electron transport rate was higher. Higher post-silking dry matter assimilation in the SG hybrid was compensated for by higher dry matter remobilisation in the non-SG. Higher kernel number per plant in the non-SG hybrid with no ability to compensate for by higher kernel weight in the SG, resulted in slightly higher yields in the non-SG under drought. A water conservation strategy associated with a shallower root system could be linked to the SG trait, promoting faster water depletion at upper soil levels and reduced stomatal conductance at the leaf level but with no conclusive yield advantage.

In C3 cereals such as wheat and barley, grain filling was traditionally explained as being sustained by assimilates from concurrent leaf photosynthesis and remobilization from the stem. In recent decades, a role for ear photosynthesis as a contributor to grain filling has emerged. This review analyzes several aspects of this topic: (i) methodological approaches for estimation of ear photosynthetic contribution to grain filling; (ii) the existence of genetic variability in the contribution of the ear, and evidence of genetic gains in the past; (iii) the controversy of the existence of C4 metabolism in the ear; (iv) the response of ear photosynthesis to water deficit; and (v) morphological and physiological traits possibly related to ear temperature and thermal balance of the ear. The main conclusions are: (i) there are a number of methodologies to quantify ear photosynthetic activity (e.g. gas exchange and chlorophyll fluorescence) and the contribution of the ear to grain filling (individual ear shading, ear emergence in shaded canopies, and isotope composition); (ii) the contribution of ear photosynthesis seems to have increased in modern wheat germplasm; (iii) the contribution of the ear to grain filling increases under resource-limitation (water deficit, defoliation, or pathogen infection); (iv) there is genetic variability in the contribution of the ear in wheat, opening up the possibility to use this trait to ameliorate grain yield; (v) current evidence supports the existence of C3 metabolism rather than C4 metabolism; (vi) the ear is a ‘dehydration avoider organ’ under drought; and (vii) thermal balance in the ear is a relevant issue to explore, and more research is needed to clarify the underlying morphological and physiological traits.

Small grain cereals such as wheat, rice and barley are among the most important crops worldwide. Any attempt to increase crop productivity and stability through breeding implies developing new strategies for plant phenotyping, including defining ideotype attributes for selection. Recently, the role of non-foliar photosynthetic organs, particularly the inflorescences, has received increasing attention. For example, ear photosynthesis has been reported to be a major contributor to grain filling in wheat and barley under stress and good agronomic conditions. This review provides an overview of the particular characteristics of the ear that makes this photosynthetic organ better adapted to grain filling than the flag leaf and revises potential metabolic and molecular traits that merit further research as targets for cereal improvement. Currently, the absence of high-throughput phenotyping methods limits the inclusion of ear photosynthesis in the breeding agenda. In this regard, a number of different approaches are presented.

In crop studies, shading is a common approach for reducing source activity (i.e. assimilates produced by photosynthesis); however, if source reduction is not directly proportional to the degree of light attenuation, estimations could be flawed. We analysed the effects of shading at three levels on light transmittance, air and canopy temperatures, and actual photosynthetic activity at different times of day, and on grain weight in a modern genotype of bread wheat (Triticum aestivum L.). The treatments were imposed 10 days after anthesis and comprised three levels of shading with meshes: 80%, 50% and 10% transmittance during midday. The decrease in midday photosynthetic activity (electron transport rate, ETR; and CO2 assimilation, An) in the shading treatments compared with control plants did not fit a 1:1 relationship with light transmittance of the meshes; for example, the reduction in ETR and An was

Green parts of the ear (e.g. bracts, awns) are sources of photosynthates in C3 cereals. However, the contribution of ear photosynthesis to grain filling in bread wheat (Triticum aestivum L.) is less known compared with other sources of assimilates (e.g. flag leaf photosynthesis, C stem retranslocation). The main objectives of this work were: (i) to analyze if empirical breeding has changed the contribution of ear photosynthesis to grain filling through a retrospective study of cultivars of bread wheat released in Argentina between 1920 and 2008 and (ii) to study if the introgression of dwarfism alleles has changed the contribution of ear photosynthesis to grain filling using near-isogenic lines of a cultivar of bread wheat differing in absence/presence in Rht alleles. In both experiments we covered ears with aluminum foil to reduce photosynthesis, in intact and defoliated plants. Grain weight per ear was measured and the ear contribution to grain filling was calculated as: [(GWear of non-shaded ear − GWear of shaded ear) × 100/GWear of non-shaded ear], where GWear is total grain weight per ear. The contribution of ear photosynthesis increased in modern cultivars, more in defoliated than intact plants. Ear contribution was negatively correlated with plant height (r2 = 0.62 and 0.71 in intact and defoliated plants, respectively) and especially with stem dry weight (r2 = 0.75 and 0.80 in intact and defoliated plants respectively). Dry matter translocation efficiency (DMTe) did not change consistently with year of release of the cultivars. The contribution of preanthesis stem assimilates to grain yield (CPA, i.e. the ratio of stem mobilized reserves to GWear) clearly decreased in more modern cultivars. The correlation of ear contribution with awn size was rather low in this retrospective study. In near-isogenic lines differing in Rht alleles, the contribution of the ear was higher when dwarfism alleles were present, mainly in non-defoliated plants. However, there was no consistent relationship between stem weight of the near-isogenic lines and their respective contributions of ear photosynthesis, suggesting that other factors (beyond the Rht alleles) are involved in the increase of ear contribution in the retrospective study.

The contribution of ear photosynthesis to grain filling in wheat (Triticum aestivum L.) is not well known. The main objective of this work was to evaluate this contribution through three different experimental approaches: (1) ear photosynthesis was reduced by removing awns or shading the ears (in combination with a defoliation treatment), (2) grain weight per ear was compared in an ‘all shaded’ crop versus plants where only the vegetative parts were shaded (‘ear emerging’), and (3) ear photosynthesis was reduced with DCMU (3-(3,4-dichlorophenyl)-1,1-dimethylurea), a specific inhibitor of photosystem II. In field experiments in La Plata (Argentina), cultivars Klein Escudo and BioINTA 3000 were subjected to awn removal and ear shading treatments, with or without severe defoliation, and to ‘all canopy shaded’ versus ‘ear emerging’ treatments. Although the estimated contribution of ear photosynthesis to grain yield differed depending on the experimental approach used (from about 12–42%), in general cv. Klein Escudo (with the largest awns) showed a greater contribution of ear photosynthesis. In both cultivars, the percentage contribution of ear photosynthesis was larger for defoliated plants. The magnitude of this contribution was positively associated with the length of awns of each cultivar. Awn removal had a non-significant (p ≤ 0.05) effect on grain weight per ear (GWear) in the short-awned cv. BioINTA 3000. In cv. Klein Escudo, the effects of de-awning were larger, reaching values of ca. 15 and 19% decrease of GWear in non-defoliated and defoliated plants respectively. In both cultivars, grains in a distal position within the spikelet (G3) were most affected by the decrease of the photosynthetic source. The photosynthetic rate of the ear was higher in cv. Klein Escudo than in cv. BioINTA 3000. We analyzed the contribution of remobilization of pre anthesis assimilates (stem plus sheaths). Comparing both cultivars, dry matter translocation from the stem showed the opposite pattern to the contribution of ear photosynthesis, i.e. it was higher in cv. BioINTA 3000 than in Klein Escudo. In Bordenave, Argentina, ear shading treatments revealed an important contribution of ear photosynthesis to grain yield in cv. Huenpan (a long awned, water-deficit tolerant genotype), either under water deficit or irrigation, whereas in cv. Baguette Premiun 11 (short awns) there was apparently no effect of ear photosynthesis on yield. Finally, inhibition of ear photosynthesis with DCMU reduced grain weight per ear to a similar extent as ear shading. Ear photosynthesis might represent a “buffer” to maintain grain yield under source limitations (e.g. defoliation, water stress conditions), and could have an important role even without stress, because an incipient ‘source’ limitation might be emerging in modern cultivars of bread wheat.

The awns of the ear have been reported as important sources of assimilates in wheat. However, their actual importance in grain filling is not well known. We examined several aspects of the awns and their role in the grain filling, and the objectives of this work were: (i) to study their photosynthetic activity during grain filling, comparing flag leaf, ear body and awns in two cultivars differing in ear contribution to grain filling, (ii) to estimate the influence of awns on the temperature and the hydraulic conductance of the ear, (iii) to analyze the response of the flag leaf versus awns to water deficit, particularly regarding relative water content and photosynthetic activity, (iv) to analyze if there is a correlation between awn size versus ear and awn contribution to the grain filling in six modern Argentinean cultivars, and, (v) to explore if awn size and awn contribution have changed during the past century, analyzing old and modern Argentinean cultivars. In this paper we show that: (1) during late grain filling, ear parts (particularly the body of the ear) maintained a higher photosynthetic activity than the flag leaf, (2) awns showed some ‘tolerance’ to water deficit, maintaining a higher relative water content and electron transport rate than the flag leaf under drought, (3) awns increase water conductance of the ear (particularly in the cv. K. Escudo), decrease the ear temperature during the morning, but increase ear temperature after midday (4) there is a moderately positive relationship between awn size and contribution of the ear to grain filling, and (5) in the retrospective analysis we did not find a clear tendency in awn size and awn contribution to grain filling along the years. However, in general terms modern cultivars seem to have a higher awn contribution (and higher awns size) than old ones. In summary, awns may have positive (increased photosynthetic area and activity, tolerance to drought) and negative (increased ear temperature) effects on grain filling, and their final impact on yield will depend on the interaction between these effects and prevailing environmental conditions. Fil: Maydup, Maria Luján. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Instituto de Fisiología Vegetal. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Instituto de Fisiología Vegetal; Argentina Fil: Antonietta, Mariana. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Instituto de Fisiología Vegetal. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Instituto de Fisiología Vegetal; Argentina Fil: Graciano, Corina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Instituto de Fisiología Vegetal. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Instituto de Fisiología Vegetal; Argentina Fil: Guiamet, Juan José. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Instituto de Fisiología Vegetal. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Instituto de Fisiología Vegetal; Argentina Fil: Tambussi, Eduardo Alberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Instituto de Fisiología Vegetal. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Instituto de Fisiología Vegetal; Argentina

Las partes verdes de la espiga (e.g. brácteas, aristas) son fuente de asimilados en cereales C3. Sin embargo, la contribución de la fotosíntesis de la espiga en trigo pan (Triticum aestivum L.) es menos conocida que el resto de las fuentes de asimilados (v.g. fotosíntesis de la hoja bandera, retranslocados del tallo). En trigo es ampliamente aceptado que la limitación por fuente no está presente en cultivares antiguos (Kruk et al. 1997; Borrás et al. 2004). Sin embargo, en germoplasma moderno, la limitación por fuente podría estar emergiendo (e.g. Álvaro 2008a). En este contexto los asimilados fijados por los órganos verdes de la espiga (v.g. glumas y aristas) podrían ser más relevante para sostener el llenado de granos. Capítulo 2: El principal objetivo de este capítulo fue evaluar la contribución de la espiga mediante tres aproximaciones experimentales diferentes: (1) reduciendo la fotosíntesis de la espiga por remoción de las aristas o sombreo de las espigas (combinados con un tratamiento de desfoliado) y (2) comparando el peso de granos por espiga, en cultivos completamente sombreados versus plantas donde únicamente se les sombreó la parte vegetativa (‘espigas emergiendo’) y (3) reduciendo la fotosíntesis de la espiga con DCMU (3-(3,4-dichlorophenyl)-1,1-dimethylurea) un inhibidor específico del foositema II. En experimentos a campo en La Plata (Argentina), los cultivares Klein Escudo and BioInta 3000, fueron sujetos a los tratamientos de remoción de las aristas, sombreado de espigas, con y sin desfoliación, y a los tratamientos de ‘sombreo total’ vs. el de ‘espigas emergiendo’. Aunque la estimación de la contribución de la fotosíntesis de la espiga difirió, dependiendo de la aproximación experimental usada (desde 12 a 42%), el cv. Escudo (con mayor proporción de aristas) mostró una mayor contribución en la fotosíntesis de la espiga. En ambos cultivares la mayor contribución de la espiga se encontró en plantas desfoliadas. La magnitud de esta contribución se relacionó positivamente con el largo de las aristas de cada cultivar. La remoción de las aristas no tuvo un efecto significativo (p≤ 0.05) sobre el peso de grano en el cultivar de aristas cortas (BioInta 3000). El efecto del desaristado fue mayor en el cv. Escudo, alcanzando valores de reducción del peso de granos por espiga de ca. 15 y 19% en plantas no desfoliadas y desfoliadas respectivamente. En los dos cultivares, la posición de granos más afectada por la disminución de la fuente fotosintética fue la de los granos distales dentro de la espiguilla. La tasa fotosintética de la espiga (medida por IRGA) fue mayor en el cv Escudo que en el cv. BioInta 3000. También se analizó la contribución de la remobilización de los asimilados acumulados en pre-antesis (tallo + vainas). Comparando ambos cultivares, la materia seca translocada desde los tallos mostró el patrón opuesto a la contribución de la fotosíntesis de la espiga, i.e. fue mayor en el cv. BioInta 3000 que en el cv. Escudo. En Bordenave, Argentina, el sombreo de las espigas reveló una importante contribución de la fotosíntesis de la espiga al llenado de granos en el cv. Huenpan (un genotipo de aristas largas, y tolerante al déficit hídrico), tanto bajo condiciones de riego como de secano, mientras que en el cv. Baguette Premiun 11 (de aristas cortas) aparentemente no hubo efecto de la fotosíntesis de la espiga sobre el peso de granos. Por último, la reducción de la fotosíntesis de la espiga con DCMU redujo el peso de granos por espiga en proporción similar al sombreo de las espigas por ‘encapuchado’. Capítulo 3: El principal objetivo de este capítulo fue: (1) analizar si la mejora empírica ha cambiado la contribución de la fotosíntesis de la espiga al llenado de granos, a través de un estudio retrospectivo con cultivares de trigo pan liberados en Argentina entre 1920 y 2008, y (2) estudiar si la introgresión de los alelos de enanismo han cambiado la contribución de la fotosíntesis de la espiga al llenado de granos, usando líneas isogénicas de un cultivar de trigo pan difiriendo en la presencia/ausencia en los alelos Rht. En los dos experimentos se sombrearon espigas con papel aluminio para reducir la fotosíntesis, tanto en plantas intactas como desfoliadas. La contribución de la fotosíntesis de la espiga se incrementó en los cultivares modernos, y esta contribución fue mayor en las plantas desfoliadas que en las intactas. La contribución de la espiga se correlacionó negativamente con la altura de planta (r2= 0.62 y 0.71 en plantas intactas y desfoliadas respectivamente) y especialmente con el peso de tallo (r2= 0.75 y 0.80 en plantas intactas y desfoliadas respectivamente). La eficiencia de materia seca retranslocada (es decir, la relación entre la materia seca retranslocada y el peso de tallo en antesis; DMTe) no se modificó considerablemente con el año de liberación de los cultivares. La contribución de asimilados pre-antesis al granos (CPA, i.e. la relación de reservas del tallo movilizadas y el peso de granos por espiga) claramente disminuyó en los cultivares más modernos. La correlación entre la contribución de la espiga y el tamaño de aristas fue despreciable en este estudio. En las líneas isogénicas diferenciadas en los alelos de enanismo Rht, la contribución de la espiga fue mayor cuando los alelos estuvieron presentes, principalmente en plantas no desfoliadas. Sin embargo, no hubo una relación consistente entre el peso de tallo y las respectivas contribuciones de las espigas, sugiriendo que otros factores (además de los alelos de enanismo) están involucrados en el incremento de la contribución de la espiga en el estudio retrospectivo. Capítulo 4: Los objetivos de este capítulo fueron: (1) estudiar la actividad fotosintética (tasa de transporte de electrones o ETR, indirectamente por fluorescencia de la clorofila) durante el llenado de granos, de la hoja bandera, el cuerpo de la espiga y de las aristas en dos cultivares con diferente contribución de la espiga al llenado de granos; (2) analizar la respuesta diferencial de la hoja bandera vs. las aristas al déficit hídrico mediante la medición de variables de estado hídrico (contenido relativo de agua, potencial hídrico) y actividad fotosintética (tasa de transporte de electrones o ETR); (3) estimar la influencia de la presencia/ausencia de aristas en la temperatura del cuerpo de la espiga; (4) analizar si existe correlación entre el tamaño de las aristas vs. la contribución de las aristas/espiga al llenado de granos en seis variedades modernas de cultivares argentinos con distinto tamaño de aristas; (5) explorar si el tamaño de aristas han cambiado durante la última centuria (estudio retrospectivo). En este capítulo hemos reportado que (1) durante el final del llenado de los granos, las partes de la espiga (aristas y particularmente el cuerpo de la espiga, es decir glumas y lemmas) podrían tener mayor actividad fotosintética que la hoja bandera; (2) la presencia de aristas podría tener un efecto levemente favorable (i.e. enfriando) sobre la temperatura de la espiga durante la mañana, pero deletéreo (i.e. aumentando la temperatura de la espiga) después del mediodía; (3) una característica positiva de las aristas es una cierta “tolerancia” al déficit hídrico, manteniendo un contenido relativo de agua y una ETR mayor que la hoja bandera bajo sequía; (4) encontramos una correlación positiva (moderada bajo condiciones de secano) entre el tamaño de las aristas y la contribución de la espiga durante el llenado de granos y (5) en el análisis retrospectivo de los cultivares de trigo pan de Argentina no encontramos una clara tendencia a lo largo de la mejora del tamaño de las aristas ni de la contribución de las aristas durante el llenado de granos., Facultad de Ciencias Naturales y Museo

Fast development of seedling leaf area is a relevant trait in order to increase early resource acquisition. The use of semi-dwarf genotypes of wheat has decreased early vigour of modern cultivars. We studied early vigour of 20 cultivars cropped in Argentina, and our main objectives were: (i) to analyse the genotypic variability in early vigour; (ii) to study morphological traits that can be good indicators of early vigour, such as seed mass, leaf width, and specific leaf area; and (iii) to determine whether increased dry mass allocation to roots impacts negatively on early vigour. Experiments with non-size-selected and size-selected seeds were carried out in a greenhouse. A field trial was also conducted in order to test the reliability of the greenhouse results. Seeds mass was the main parameter related to early vigour. However, results from the experiment with seeds selected by size (45-50mg seed-1) showed that seed mass per se only partially explains early vigour, since a significant coefficient of determination was observed between the seedling leaf area of each cultivar in both experiments (i.e. with randomly chosen or size-selected seeds). We observed a high coefficient of determination between net assimilation rate and changes in the ranking of early vigour of the cultivars with time after transplant. Root biomass was positively correlated with leaf area, indicating that the traits were not mutually exclusive. We built simple models by multiple regression to predict early vigour, including some parameters that were easy to measure. Seed mass and leaf width taken together showed better fit than seed mass or leaf width alone. We found a significant coefficient of determination between early vigour in greenhouse and field experiments; thus, screening for early vigour under semi-controlled conditions may be feasible. Fil: Maydup, Maria Luján. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Instituto de Fisiología Vegetal. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Instituto de Fisiología Vegetal; Argentina Fil: Graciano, Corina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Instituto de Fisiología Vegetal. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Instituto de Fisiología Vegetal; Argentina Fil: Guiamet, Juan José. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Instituto de Fisiología Vegetal. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Instituto de Fisiología Vegetal; Argentina Fil: Tambussi, Eduardo Alberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Instituto de Fisiología Vegetal. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Instituto de Fisiología Vegetal; Argentina