María E. Otegui

Growth, water use, and kernel abortion of maize subjected to drought at silking

Abstract Maize (Zea mays L.) grain yield is particularly sensitive to water deficits that coincide with the tasseling-silking period, causing marked reductions in grain number. More knowledge about crop responses to water supply is required, however, to explain the causes of kernel number reductions under the mild stresses characteristic of humid regions. The objectives of this study were to: (i) quantify crop evapotranspiration, Ec, and its relationship with shoot biomass production, grain yield, and kernel number; and (ii) determine the impact on final kernel number of supplying fresh pollen to silks whose appearance is delayed by water deficits at silking. Field experiments were conducted at Balcarce (37°45′S, 130 m) during 1988/89 and 1989/90 with two sowing dates (6 weeks apart) to provide differences in evaporative demand. Plastic covers were placed on the ground of water-deficit plots to generate a 40-day period of lowered water supply bracketing silking. Control plots received rain plus additional furrow irrigation in order to keep the ratio between crop (c) and potential (Ep) Penman evapotranspiration greater than 0.9. Plant water status indicators revealed differences between treatments, but failed to reflect soil water status. Water deficit reduced plant height, maximum leaf area index, and shoot biomass. Shoot biomass accumulation was correlated with Ec, but higher water-use efficiencies (WUE) were found for the water-stress treatments. Grain yield was correlated to kernels m−2 (r = 0.88; 6 d.f.), and both grain yield and kernels m−2 were related to Ec during the treatment period, resulting in reductions of 4.7 grains m−2 and 17.7 kg ha−1 for each mm reduction in Ec. The number of kernels per ear did not improve when fresh pollen was applied to late appearing silks, suggesting that ovaries which failed to expose their silks synchronously with pollen shedding were deleteriously affected by water stress.

Grain yield components in maize: I. Ear growth and kernel set

Abstract Maize (Zea mays, L) grain yield is correlated with kernel number (KN), but uncertainty exists about the extension of the critical period for kernel set. In this work, the actual period of active ear elongation was determined and defined in thermal units (°C day=degree days). Kernel number was related to the amount of intercepted photosynthetically active radiation (IPAR) during this period. Field experiments were conducted in France (48°51′N) and Argentina (32 to 37°S). Different sowing dates and plant populations were used to vary the amount of IPAR plant−1. Hybrids of different cycle length (FAO maturity ranking between 300 and 630) were tested. Ear length was measured weekly after axillary bud differentiation, and values were normalized to final ear length. A significant linear model (r2=0.92, n=51) was fitted, which indicated that: (i) Active ear elongation took place between −227 and 100°C day from silking; (ii) ca. 41% of final ear length was reached at silking; (iii) Ear elongation rate increased after silking; and (iv) The duration of this period varied among sites and treatments. Kernel number plant−1 was significantly related to the IPAR plant−1 during the critical period (r2=0.70, n=81), but a significant (P

Maize Leaves Turn Away from Neighbors

In commercial crops, maize (Zea mays) plants are typically grown at a larger distance between rows (70 cm) than within the same row (16–23 cm). This rectangular arrangement creates a heterogeneous environment in which the plants receive higher red light (R) to far-red light (FR) ratios from the interrow spaces. In field crops, the hybrid Dekalb 696 (DK696) showed an increased proportion of leaves toward interrow spaces, whereas the experimental hybrid 980 (Exp980) retained random leaf orientation. Mirrors reflecting FR were placed close to isolated plants to simulate the presence of neighbors in the field. In addition, localized FR was applied to target leaves in a growth chamber. During their expansion, the leaves of DK696 turned away from the low R to FR ratio signals, whereas Exp980 leaves remained unaffected. On the contrary, tillering was reduced and plant height was increased by low R to FR ratios in Exp980 but not in DK696. Isolated plants preconditioned with low R/FR-simulating neighbors in a North-South row showed reduced mutual shading among leaves when the plants were actually grouped in North-South rows. These observations contradict the current view that phytochrome-mediated responses to low R/FR are a relic from wild conditions, detrimental for crop yield.

Leaf area, light interception, and crop development in maize

Few models used to estimate the fraction of photosynthetically active radiation intercepted by maize crops (fIPAR) as a function of leaf area index (LAI) account for genotype differences and ontogenetic stage. In this study, the development of the fIPAR/GLAI relationship of three maize hybrids having contrasting plant type and grown in different environmental conditions was characterized. Three field experiments were conducted at Rojas (34°08′S), Argentina, on a silty clay loam soil, without nutrient restriction. The effects of sowing date and water regime were tested. Plant density was always 7 plants m−2. The fIPAR was calculated from measurements above and below the canopy, before and after maximum GLAI was attained. Hybrids differed significantly (P 0.77; n ≥ 15) the fIPAR/GLAI relationship for pre-maximum GLAI data of all hybrids. Nevertheless, maximum fIPAR was always below values quoted in the literature (< 0.90), and differences were detected among hybrids in the attenuation coefficient (k) that were probably related to leaf angle and leaf area. For hybrids with a similar leaf angle at all leaf positions no difference existed in the fIPAR/GLAI relationship along the cycle (pre- and post-maximum GLAI). The relationship changed after tasselling for the hybrid with erect upper leaves. Ignoring such differences could be misleading when fIPAR/GLAI models are used to estimate canopy photosynthesis and hence biomass production, radiation-use efficiency, and kernel set of specific hybrids and growth stages.

Grain yield components in maize: II. Postsilking growth and kernel weight

Maize kernel weight (KW) results from kernel growth during two stages of grain filling, the lag phase (formative period) and the effective grain-filling phase. Environmental conditions may affect kernel biomass accumulation in each phase. This work analyzed: (1) changes in duration and rate of kernel growth on a thermal time (°C day) basis; and (2) KW response to postsilking biomass production kernel−1 (source:sink ratio). Sowing date, plant population, and nitrogen fertilization experiments were conducted in France and Argentina to induce changes in assimilate availability per kernel. Hybrids of different KW were tested. Hybrids differed in the duration of the lag phase, which determined kernel growth rate during the effective grain-filling period for hybrids with similar grain-filling duration (ca. 745°C day). Environments with low air temperature ( 300 mg) with reduced kernel number (2800 to 4000 kernels m−2). For the former, grain yield increments should not be based on increased kernel number but on increased biomass production.

Kernel weight dependence upon plant growth at different grain-filling stages in maize and sorghum

In the present study we tested how assimilate availability per kernel at different grain-filling stages may affect maize (Zea mays L.) and sorghum (Sorghum bicolor L. Moench) individual kernel weight (KW). These two species have shown a contrasting KW response to increased assimilate availability at similar seed developmental stages. Plant growth rate (PGR) per kernel was used to estimate the assimilate availability per kernel at two stages: around the early grain-filling period when kernel number per plant is also being established, and around the effective grain-filling period. We tested 3 commercial genotypes from each species, and modified the PGR by thinning or shading the stand at different developmental stages. In both species, each genotype showed a particular relationship between PGR around flowering and kernel number, which gave a range of responses in the PGR per kernel set around flowering. Final KW always increased whenever PGR per kernel around flowering was enhanced. Only sorghum showed a consistent KW increase when PGR per kernel during the effective grain-filling period was enhanced. Results confirmed that increasing assimilate availability per kernel will affect maize kernel size only if the potential set early in development is altered. Most important, we showed that linking specific KW sensibility across species at different seed developmental stages using a simple estimate of assimilate availability per seed (i.e. PGR per kernel) at each grain-filling stage helped explain most of the explored genotypic and environmental variability in final kernel size.

Silk Elongation in Maize

In maize (Zea mays L.), the gradient in floret development and silk length along the ear at silking determines a time lag between early- and late-appearing silks, which results in pollination asynchrony between them. This asynchrony is partially responsible of reduced kernel set at the ear tip, and hybrids differ in this trait. The objective of this work was to analyze the pattern of floret and silk differentiation and elongation at different spikelet positions (S n ) along the apical ear of two hybrids of contrasting ear size (DEA ≅ 500 spikelets ear -1 ; DK696 ≅ 800 spikelets ear -1 ). At silking, both hybrids had reached approximately the same proportion of final ear length (about 44%), but DK696 had differentiated a greater number of spikelets row -1 (46 spikelets) than DEA (33 spikelets). Silk initiation rate was always faster than spikelet initiation rate, and silk extension dynamics was similar for all spikelet positions. Silks from the base of the ear were always longer than those from the tip (S 25 in DEA or S 35 in DK696). Before pollination, silks experienced an early phase of exponential elongation followed by a phase of linear growth. A drastic reduction in elongation rate followed silk emergence, which did not occur when ears were bagged and pollination was prevented. Convergence in silking among spikelets along the ear could be attained by (i) synchronous silk initiation among spikelet positions, followed by a similar pattetn of silk elongation in all florets (hybrid DEA), or (ii) increased silk elongation rate in apical florets (hybrid DK696).

Water deficit stress tolerance in maize conferred by expression of an isopentenyltransferase (IPT) gene driven by a stress- and maturation-induced promoter

Senescence can be delayed in transgenic plants overexpressing the enzyme isopentenyltransferase (IPT) due to stress-induced increased levels of endogenous cytokinins. This trait leads to sustained photosynthetic activity and improved tolerance to abiotic stress. The aim of this study was to generate and characterize transgenic plants of maize (Zea mays L.) transformed with the IPT gene sequence under the regulation of SARK promoter (protein kinase receptor-associated senescence). Three independent transgenic events and their segregating null controls were evaluated in two watering regimes (WW: well watered; WD: water deficit) imposed for two weeks around anthesis. Our results show that the WD treatment induced IPT expression with the concomitant increase in cytokinin levels, which prolonged the persistence of total green leaf area, and maintained normal photosynthetic rate and stomatal conductance. These trends were accompanied by a minor decrease in number of grains per plant, individual grain weight and plant grain yield as compared to WW plants. Plants expressing the IPT gene under WD had PGR, anthesis and silking dates and biomass levels similar to WW plants. Our results demonstrate that expression of the IPT gene under the regulation of the SARK promoter helps improve productivity under WD conditions in C4 plants like maize.

A role for LAX2 in regulating xylem development and lateral-vein symmetry in the leaf

Background and Aims The symmetry of venation patterning in leaves is highly conserved within a plant species. Auxins are involved in this process and also in xylem vasculature development. Studying transgenic Arabidopsis plants ectopically expressing the sunflower transcription factor HaHB4, it was observed that there was a significant lateral-vein asymmetry in leaves and in xylem formation compared to wild type plants. To unravel the molecular mechanisms behind this phenotype, genes differentially expressed in these plants and related to auxin influx were investigated. Methods Candidate genes responsible for the observed phenotypes were selected using a co-expression analysis. Single and multiple mutants in auxin influx carriers were characterized by morphological, physiological and molecular techniques. The analysis was further complemented by restoring the wild type (WT) phenotype by mutant complementation studies and using transgenic soybean plants ectopically expressing HaHB4 . Key Results LAX2 , down-regulated in HaHB4 transgenic plants, was bioinformatically chosen as a candidate gene. The quadruple mutant aux1 lax1 lax2 lax3 and the single mutants, except lax1, presented an enhanced asymmetry in venation patterning. Additionally, the xylem vasculature of the lax2 mutant and the HaHB4 -expressing plants differed from the WT vasculature, including increased xylem length and number of xylem cell rows. Complementation of the lax2 mutant with the LAX2 gene restored both lateral-vein symmetry and xylem/stem area ratio in the stem, showing that auxin homeostasis is required to achieve normal vascular development. Interestingly, soybean plants ectopically expressing HaHB4 also showed an increased asymmetry in the venation patterning, accompanied by the repression of several GmLAX genes. Conclusions Auxin influx carriers have a significant role in leaf venation pattering in leaves and, in particular, LAX2 is required for normal xylem development, probablt controlling auxin homeostasis.

Water deficit and impaired pegging effects on peanut seed yield: links with water and photosynthetically active radiation use efficiencies

Peanut (Arachis hypogaea L.) production is frequently affected by unpredictable events of water deficit during pod set, which modulate water use, water use efficiency for biomass production (WUEB), and biomass partitioning to seeds. We studied the effects of drought-induced impaired pegging on WUEB and the link between WUEB and photosynthetically active radiation use efficiency (PAR-UE). Field experiments were conducted that combined: two cultivars of contrasting pegging capacity (ASEM > Florman), two water regimes (irrigated and water stress) and different sowing dates. WUEB ranged between 6.1 and 6.7 g kPa/mm for irrigated plots, and between 2.9 and 7.1 g kPa/mm for water-stressed plots. WUE for pod production showed similar trends, but was larger for ASEM than for Florman because of higher biomass allocation to pods and pegging capacity of the former. The relationship between standardised values of WUEB and PAR-UE varied linearly for the post-R6 period, but fitted models differed between water regimes. This difference was attributed to the relative importance of stomata control on gas exchange (direct effects of water deficit) respect to feedback effects on photosynthesis caused by reproductive sink size (indirect effects of water deficit). Variation in post-R6 PAR-UE could be linked exclusively to the latter, but variation registered in WUEB acknowledged both controls.