Stress-induced expression of IPT gene in transgenic wheat reduces grain yield penalty under drought.

BACKGROUND: The heterologous expression of isopentenyl transferase (IPT) under the transcriptional control of the senescence-associated receptor-like kinase (SARK) promoter delayed cellular senescence and, through it, increased drought tolerance in plants. To evaluate the effect of pSARK::IPT expression in bread wheat, six independent transgenic events were obtained through the biolistic method and evaluated transgene expression, phenology, grain yield and physiological biomass components in plants grown under both drought and well-irrigating conditions. Experiments were performed at different levels: (i) pots and (ii) microplots inside a biosafety greenhouse, as well as under (iii) field conditions. RESULTS: Two transgenic events, called TR1 and TR4, outperformed the wild-type control under drought conditions. Transgenic plants showed higher yield under both greenhouse and field conditions, which was positively correlated to grain number (given by more spikes and grains per spike) than wild type. Interestingly, this yield advantage of the transgenic events was observed under both drought and well-watered conditions. CONCLUSIONS: The results obtained allow us to conclude that the SARK promoter-regulated expression of the IPT gene in bread wheat not only reduced the yield penalty produced by water stress but also led to improved productivity under well-watered conditions.

Soybean fruit development and set at the node level under combined photoperiod and radiation conditions.

In soybean, long days during post-flowering increase seed number. This positive photoperiodic effect on seed number has been previously associated with increments in the amount of radiation accumulated during the crop cycle because long days extend the duration of the crop cycle. However, evidence of intra-nodal processes independent of the availability of assimilates suggests that photoperiodic effects at the node level might also contribute to pod set. This work aims to identify the main mechanisms responsible for the increase in pod number per node in response to long days; including the dynamics of flowering, pod development, growth and set at the node level. Long days increased pods per node on the main stems, by increasing pods on lateral racemes (usually dominated positions) at some main stem nodes. Long days lengthened the flowering period and thereby increased the number of opened flowers on lateral racemes. The flowering period was prolonged under long days because effective seed filling was delayed on primary racemes (dominant positions). Long days also delayed the development of flowers into pods with filling seeds, delaying the initiation of pod elongation without modifying pod elongation rate. The embryo development matched the external pod length irrespective of the pod's chronological age. These results suggest that long days during post-flowering enhance pod number per node through a relief of the competition between pods of different hierarchy within the node. The photoperiodic effect on the development of dominant pods, delaying their elongation and therefore postponing their active growth, extends flowering and allows pod set at positions that are usually dominated.

High night temperatures during grain number determination reduce wheat and barley grain yield: a field study.

Warm nights are a widespread predicted feature of climate change. This study investigated the impact of high night temperatures during the critical period for grain yield determination in wheat and barley crops under field conditions, assessing the effects on development, growth and partitioning crop-level processes driving grain number per unit area (GN). Experiments combined: (i) two contrasting radiation and temperature environments: late sowing in 2011 and early sowing in 2013, (ii) two well-adapted crops with similar phenology: bread wheat and two-row malting barley and (iii) two temperature regimes: ambient and high night temperatures. The night temperature increase (ca. 3.9 °C in both crops and growing seasons) was achieved using purpose-built heating chambers placed on the crop at 19:000 hours and removed at 7:00 hours every day from the third detectable stem node to 10 days post-flowering. Across growing seasons and crops, the average minimum temperature during the critical period ranged from 11.2 to 17.2 °C. Wheat and barley grain yield were similarly reduced under warm nights (ca. 7% °C(-1) ), due to GN reductions (ca. 6% °C(-1) ) linked to a lower number of spikes per m(2) . An accelerated development under high night temperatures led to a shorter critical period duration, reducing solar radiation capture with negative consequences for biomass production, GN and therefore, grain yield. The information generated could be used as a starting point to design management and/or breeding strategies to improve crop adaptation facing climate change.

More fertile florets and grains per spike can be achieved at higher temperature in wheat lines with high spike biomass and sugar content at booting.

An understanding of processes regulating wheat floret and grain number at higher temperatures is required to better exploit genetic variation. In this study we tested the hypothesis that at higher temperatures, a reduction in floret fertility is associated with a decrease in soluble sugars and this response is exacerbated in genotypes low in water soluble carbohydrates (WSC). Four recombinant inbred lines contrasting for stem WSC were grown at 20/10°C and 11h photoperiod until terminal spikelet, and then continued in a factorial combination of 20/10°C or 28/14°C with 11h or 16h photoperiod until anthesis. Across environments, High WSC lines had more grains per spike associated with more florets per spike. The number of fertile florets was associated with spike biomass at booting and, by extension, with glucose amount, both higher in High WSC lines. At booting, High WSC lines had higher fixed 13C and higher levels of expression of genes involved in photosynthesis and sucrose transport and lower in sucrose degradation compared with Low WSC lines. At higher temperature, the intrinsic rate of floret development rate before booting was slower in High WSC lines. Grain set declined with the intrinsic rate of floret development before booting, with an advantage for High WSC lines at 28/14°C and 16h. Genotypic and environmental action on floret fertility and grain set was summarised in a model.

Wheat floret survival as related to pre-anthesis spike growth.

Further improvements to wheat yield potential will be essential to meet future food demand. As yield is related to the number of fertile florets and grains, an understanding of the basis of their generation is instrumental to raising yield. Based on (i) a strong positive association between the number of fertile florets or grains and spike dry weight at anthesis; and (ii) the finding that floret death occurs when spikes grow at maximum rate, it was always assumed that floret survival depends on the growth of the spike. However, this assumption was recently questioned, suggesting that assimilates diverted to the spike do not determine the number of florets and grains and that the onset of floret death may instead be a developmental process that is not associated with spike growth. In this study, the relationships between the fate of floret primordia and spike growth from six independent experiments that included different growing conditions (greenhouse/field experiments, growing seasons, photoperiod/shading treatments during the floret primordia phase) and diverse cultivar types (winter/spring, semi-dwarf/standard-height, photoperiod sensitive/insensitive) were re-analysed together. Onset of floret death was associated with the beginning of spike growth at the maximum rate in c. 80% of the cases analysed; and the rate of floret death (the main determinant of floret survival) showed a negative quantitative relationship with spike weight at anthesis. As floret death and survival were shown to be linked to pre-anthesis spike growth, the strategy of focusing on traits associated with pre-anthesis spike growth when breeding to increase wheat yield potential further is valuable.

Changes in duration of developmental phases of durum wheat caused by breeding in Spain and Italy during the 20th century and its impact on yield.

BACKGROUND AND AIMS: Although the apical development of wheat has been widely described, studies analysing how genetic breeding over the 20th century influenced the developmental phases and its consequences on yield generation are lacking, especially for durum wheat under field conditions in Mediterranean environments. The aims of this study were to analyse the effects of breeding in Spain and Italy on crop development during the last century, to determine whether or not breeding significantly altered the developmental phases between sowing and maturity, and to evaluate the importance of each phase in determining the number of grains per spike of durum wheat (Triticum durum) cultivars representing the germplasm grown throughout the 20th century in Spain and Italy. METHODS: Eight field experiments were carried out during 4 years in two contrasting latitudes (Lleida and Granada, Spain). Plant material consisted of 24 durum wheat cultivars (12 Italian and 12 Spanish) grown throughout the 20th century in Spain and Italy. KEY RESULTS: In Spanish materials, breeding reduced the duration of the period from sowing to anthesis, placing the grain-filling period in better conditions. In those cultivars, the sub-phase sowing-terminal spikelet formation was reduced while the duration of the period from booting to anthesis was increased. The number of grains per spike increased by 23 % from old to modern cultivars, by changes in the number of grains per spikelet in both Spanish and Italian cultivars. Floral abortion from booting to anthesis diminished by 24 % from old to modern cultivars, and grain setting increased by 13 %. CONCLUSIONS: The results suggest that breeding reduced not only plant height, but also the time to anthesis. By extending the duration of the phase from booting to anthesis, which was associated with an increase in spike dry weight and grains per spike, it suggests that future increases in spike fertility could be achieved by enlarging that phase.

Autophagy regulated by day length determines the number of fertile florets in wheat.

SUMMARY: The wheat spikelet meristem differentiates into up to 12 floret primordia, but many of them fail to reach the fertile floret stage at anthesis. We combined microarray, biochemical and anatomical studies to investigate floret development in wheat plants grown in the field under short or long days (short days extended with low-fluence light) after all the spikelets had already differentiated. Long days accelerated spike and floret development and greening, and the expression of genes involved in photosynthesis, photoprotection and carbohydrate metabolism. These changes started while the spike was in the light-depleted environment created by the surrounding leaf sheaths. Cell division ceased in the tissues of distal florets, which interrupted their normal developmental progression and initiated autophagy, thus decreasing the number of fertile florets at anthesis. A massive decrease in the expression of genes involved in cell proliferation, a decrease in soluble carbohydrate levels, and an increase in the expression of genes involved in programmed cell death accompanied anatomical signs of cell death, and these effects were stronger under long days. We propose a model in which developmentally generated sugar starvation triggers floret autophagy, and long days intensify these processes due to the increased carbohydrate consumption caused by the accelerated plant development.

Photoperiod during stem elongation in wheat: is its impact on fertile floret and grain number determination similar to that of radiation?

Increasing duration of stem elongation by exposure to short photoperiod would result in higher spike dry weight at anthesis, which is positively associated with the number of fertile florets and grains in wheat. However, it is not easy to determine whether photoperiod effects on fertile florets and grains are only mediated by assimilate supply to the growing spike when spike weight variation is attained only with photoperiod treatments. The aim of this study was to determine whether photoperiod effects on number of fertile florets and grains may be direct, that is, not mediated by assimilate supply, by comparing the magnitude of photoperiod effects with those of shading the canopy. Spike dry weight at anthesis was changed through the factorial combination of different photoperiod (natural and 6 h extended photoperiod) and shading (un-shaded and 67 ± 3% shaded) treatments during stem elongation of Buck Manantial, a cultivar known for its photoperiod sensitivity in this phase. Both treatments modified spike dry weight at anthesis and the number of fertile florets and grains, independently. When duration of stem elongation was lengthened by exposure to natural photoperiod and when incident radiation was high, spike dry weight at anthesis increased by 33% (NP+0 v. NP+6) and 27% (un-shaded v. shaded), respectively. The number of fertile florets increased similarly to spike dry weight (34% NP+0 v. NP+6 and 28% un-shaded v. shaded) resulting in higher number of grains. Most photoperiod effects on the number of fertile florets and, consequently, on the number of grains, were mediated by assimilate supply to the growing spike as the same relationship between the number of fertile florets and spike dry weight at anthesis was observed for photoperiod and shading treatments (R2 = 0.99, P<0.05).

Floret development and survival in wheat plants exposed to contrasting photoperiod and radiation environments during stem elongation.

Wheat breeding has improved yield potential increasing floret survival through higher dry matter partitioning to the spikes during the stem elongation phase (from terminal spikelet initiation to anthesis). We studied survival of floret primodia in different spikelet positions along the spike in relation to dynamics of spike growth, when dynamics of dry matter partitioning to the spike was altered by photoperiod and shading treatments applied during the stem elongation phase. The cultivar Buck Manantial was exposed to (1) NP+0 un-shaded (natural photoperiod and incoming radiation of the growing season), (2) NP+0 shaded (natural photoperiod but only 33% of the incoming radiation), and (3) NP+6 un-shaded (natural photoperiod extended 6 h and natural incoming radiation). Floret survival increased, depending on spikelet position, 1.1-2.5 fold under un-shaded v. shaded treatments (both under NP+0), and 1.3-1.8 fold under NP+0 v. NP+6 treatments (both un-shaded), without any impact of treatments on the total number of initiated floret primordia. The fate of the floret primordia and its final stage at anthesis were associated with duration of floret development within the stem elongation phase (R2 = 82%, P<0.0001). Florets may be classified into three groups: (i) those that were fertile at anthesis under all treatments (mostly the two florets F1 and F2, proximal to the rachis within the spikelet), (ii) those that reached different stages at anthesis, depending on treatment, and that contributed differentially to the number of fertile florets at anthesis (mostly the florets F3, F4 and F5, positioned in the middle of the spikelet), and (iii) those that did not contribute to the number of fertile florets under any treatment (mostly the florets ≥ F6). Degeneration of florets in group (ii) was associated with spike growth at maximum rate, explaining the strong relationship observed between spike dry weight at anthesis and number of fertile florets. However, degeneration of florets in group (iii) seemed to occur before spike growth at maximum rate. Survival of florets positioned in the middle of the spikelets could be improved by increasing spike growth through manipulation of photoperiod sensitivity during stem elongation.