Nitrogen Fertilizer Effects on the Duration of the Pre-anthesis Period and Spikelet Number per Spike in Barley

The effects of increased yield and grain number per unit area in barley in response to nitrogen application are well known. However, the influence of applied nitrogen on the rates and durations of developmental phases in barley are less well understood. Our objective was to investigate the effect of applied nitrogen on the duration of pre-anthesis development in barley and the number of spikelets per spike in two barley cultivars, Franklin and Schooner, in two studies. We found no effect of nitrogen on the duration of the pre-anthesis period in Schooner, when applied to pots at a rate of 0 or 55 kg N ha^-1, or when applied in the held at 0, 40 or 160 kg N ha^-1. However, this duration was extended in Franklin in the first study by an application of 55 kg N ha^-1. Both plant biomass and grain yield at maturity were increased between 0 and 55 kg N ha^-1, and 0 and 160 kg N ha^-1. Meld increase was largely associated with an increase in the number of tillers per plant.     

小麦穗籽粒数、单粒重及单粒蛋白质含量的小穗位和粒位效应

通过系统分析2个品种与2个施氮水平的试验资料,确立了小麦穗籽粒数、单粒重、单粒蛋白质含量与蛋白质产量的小穗位和粒位效应。结果表明,不同小穗位结实籽粒数、小穗重,不同粒位粒重、单粒蛋白质含量与蛋白质产量均呈现二次曲线分布。2个品种穗部籽粒的分布存在显著差异,多粒型品种在不同施氮水平下每小穗结实粒数、单粒     

The Effect of Late-terminal Drought Stress on Yield Components of Four Barley Cultivars

Barley (Hordeum vulgare L.) is an important winter cereal crop grown in the semiarid Mediterranean, where late‐terminal drought stress during grain filling has recently become more common. The objectives of this study were to investigate the growth performance and grain yield of four barley cultivars under late‐terminal drought stress under both glasshouse and field conditions. At grain filling, four barley cultivars (Rum, ACSAD176, Athroh and Yarmouk) were exposed to three watering treatments: (1) well‐watered [soil maintained at 75 % field capacity (FC)], (2) mild drought stress at 50 % FC, (3) severe drought stress at 25 % FC in the glasshouse experiment and (1) well‐watered (irrigated once a week), (2) mild drought (irrigated once every 2 weeks), (3) severe drought (non‐irrigated; rainfed) in the field. As drought stress severity increased, gross photosynthetic rate, water potential, plant height, grain filling duration, spike number per plant, grain number per spike, 1000‐grain weight, straw yield, grain yield and harvest index decreased. In the glasshouse experiment, the six‐row barley cultivars (Rum, ACSAD176, and Athroh) had higher grain yield than the two‐row barley cultivar (Yarmouk), but the difference was not significant among the six‐row cultivars under all treatments. In the field experiment, Rum had the highest grain yield among all cultivars under the mild drought stress treatment. The two‐row cultivar (Yarmouk) had the lowest grain yield. In general, the traditional cultivar Rum had either similar or higher grain yield than the other three cultivars under all treatments. However, the yield response to drought differed between the cultivars. Those, Rum and ACSAD176, that were capable of maintaining a higher proportion of their spikes and grains per spike during drought also maintained a higher proportion of their yield compared with those in well‐watered treatment. In conclusion, cultivar differences in grain yield were related to spike number per plant and grain number per spike, but not days to heading or grain filling duration.     

野生大麦的抗旱性及其在大麦遗传改良中的应用

干旱是主要的非生物胁迫性灾害,极大地威胁到全球的粮食供应,是影响农业生产的最重要因素之一。因此,培育抗旱性植物是解决干旱危害和粮食危机的根本途径。野生大麦已经适应广泛的干旱环境并具有丰富的遗传多样性,它们的抗旱基因与抗旱相关数量性状位点(QTLs)已经确定,对改良大麦抗旱性有巨大的潜力。综述了野生大麦的抗旱性及其在大麦改良中的应用,并对大麦抗旱育种中存在的问题以及今后发展方向进行了展望。     

Effects of Fungicide on Grain Yield of Barley Grown in Different Cropping Systems

The effect of fungicides and their combination on yield of barley under different nitrogen, slurry and tillage treatments was investigated at Hohenschulen Experimental Station near Kiel, Germany in 1991–97. Various fungicide treatments (no fungicide, and treatment with stem, leaf and ear fungicides and combinations of these), two nitrogen levels (120 and 240 kg N ha⁻¹), two tillage systems (minimum and conventional tillage) and four slurry applications (no application, and autumn, spring and autumn plus spring applications) were used. On average, fungicide application increased barley yield by 1.1 t ha⁻¹. The fungicide treatments could be classified into four types: (1) fungicides against stem diseases, which slightly increased yield by 0.25 t ha⁻¹, very similar to the results for the untreated control; (2) leaf fungicides and ear fungicides applied separately, and fungicides against a combination of stem and leaf diseases, which increased yield by 1.0 t ha⁻¹ on average; (3) fungicides against a combination of ear and stem diseases, which increased the yield by 1.22 t ha⁻¹, and (4) fungicides against a combination of leaf and ear diseases and a combination of stem, leaf and ear diseases, which increased yield by 1.59 t ha⁻¹ on average. The effects of fungicide on the yield were modified by crop husbandry. It can be concluded that application of fungicides against a combination of leaf and ear diseases could increase barley yield and reduce yield variation.     

氮素养分对青稞营养功能特征及籽粒产量的影响

为确定在不同氮素条件下影响青稞生产力的关键性营养功能特征,以‘藏青2000’为试材,就不同施氮水平对青稞营养功能特征及籽粒产量的影响进行试验研究。结果表明：施氮显著提高了青稞籽粒产量、株高、相对生长率和比叶面积,而降低了单位叶面积根量和叶干物质含量,施氮水平与株高、相对生长率和比叶面积呈正相关关系,与单位叶面积根量和叶干物质含量呈负相关关系。随着施氮强度加大,产量持续提高,但增长速率下降,青稞籽粒产量与施氮水平呈二次曲线关系(y=-171.25x ²+1327.3x+10, R ²=0.958, P小于0.05)。青稞籽粒产量与株高(y=43.026x-2596.6, R ²=0.8994, P小于0.05)、相对生长率(y=7178.7x-150.66, R ²=0.5361, P小于0.05)和比叶面积(y=6.4604x+413.3, R ²=0.4153, P小于0.05)呈显著线性正相关,与叶干物质含量(y=-29.668x+4045.7, R ²=0.4488, P小于0.05)呈线性负相关关系,与单位叶面积根量(y=-2025.4x+2746.4, R ²=0.0579, P大于0.05)无明显相关性。结果表明：青稞的植株高度、相对生长率和比叶面积促进产量增长,叶干物质含量对产量有抑制作用,而单位叶面积根量对产量影响不明显。在本试验条件下,植株高度、相对生长率、比叶面积和叶干物质含量4个功能特征可在物种水平上被认为是影响青稞生产力的标记性特征。     

Spring Barley Responses to Nitrogen Fertilizer and Ethephon in Regions with a Short Crop Growing Season

Aspects of intensive management practices such as high N fertilization inputs and plant growth regulator use could potentially increase cereal yields in regions with a short crop-growing season and occasionally dry and hot weather. A field experiment was carried out for four years (1987 to 1990) at Agronomy Research Centre of McGill University, Canada to evaluate the response of spring barley ( Hordeum vulgare L.) cv. Cadette, Laurier and Leger to N rates (0, 70 and 140 kg N ha⁻¹) and ethephon (2-chloroethyl phosphonic acid) growth regulator treatment. The application of higher than conventional N level (70 kg N ha⁻¹) did not increase barley yields under dry weather conditions and when the soil N resources were high. The high levels of N did increase the grain crude protein concentration by 2 to 10 g kg⁻¹, as did ethephon treatment. The feed quality of spring barley was improved. Ethephon treatment reduced grain yield due to a severe reduction in the number of grains spike⁻¹, and/or 1000-grain weight, although the number of spikes m⁻² was increased. Thus, the overall effect of ethephon treatment on yield was inconsistent and was influenced to a large extent by the prevailing weather conditions.     

Spring Barley Responses to Row Spacing and Fungicide Triadimefon in Regions with a Short Crop-growing Season

Aspects of intensive management practices such as high-yielding cultivars, narrow-row spacings and fungicide treatments could potentially increase cereal yields in regions with a short crop-growing season and occasionally dry and hot weather. A field experiment was carried out at McGill University, Canada for three years (1987 to 1989) to test the effects of triadimefon (1-[4-chlorophenoxy]-3,3-dimethyl-1-[1H-1,2,4-triazol-1-yl]-2-butanone, or Bayleton, a trade name) fungicide (0 vs. 140 g a.i. ha⁻¹) and row spacing (10 vs. 20 cm) on the yield components, yield and other agronomic traits (spike emergence, days to maturity, leaf disease and plant height) of spring barley ( Hordeum vulgare L.) cv. Cadette, Laurier and Leger. A cultivar by row spacing interaction resulted in a 11 to 13.5 % increase in grain yield due to narrow rows for Laurier in two out of the three years and up to 16 % for Leger in one year. Over the three years grain yields were on average increased 6 to 12 % due to use of the narrower row spacing. Fungicide application to barley at the early heading stage effectively controlled leaf diseases without influence on yield components or grain yield. Our results indicate that some components of intensive management such as narrow row spacing can be applicable in regions with a short crop-growing season.     

Effects of Mild and Severe Drought Stress on Photosynthetic Efficiency in Tolerant and Susceptible Barley (Hordeum vulgare L.) Genotypes

Drought stress is one of the most important abiotic factors which adversely affect growth, metabolism and yield of crops worldwide. The objective of this study was to determine the effects of drought stress on photosynthesis in barley and examine the differential responses of photosynthetic apparatus in relatively tolerant (Yousof) and susceptible (Morocco) barley genotypes. Plants were subjected to different levels of soil water availability including control, mild and severe drought stress. In both genotypes, drought stress led to decrease in chlorophylls, β‐carotene and stomatal conductance accompanied by decrease in CO₂ assimilation rate. Significant increase in α‐tochoperol content was only observed in Yousof cultivar under drought stress. Initial slope and plateau phase of CO₂ response curve of drought‐stressed plants as well as polyphasic chlorophyll a fluorescence transient curve (OJIP test) and fast fluorescence induction kinetics were influenced by drought stress. These parameters were more affected in Morocco cultivar by drought stress compare with Yousof. Drought stress also resulted in reduction of D1 protein content in both genotypes and accelerated photoinhibition process. Based on our results, stomatal conductance is the main factor limiting photosynthesis in Yousof genotype under mild drought stress. However, in Morocco, in addition to stomatal limitation, damage to photosystem II (PSII), reduced electron transport and carboxylation efficiencies were important parts of limitation in photosynthesis. Severe drought stress resulted in structural and biochemical impairment of light‐dependent reactions as well as carboxylation process of photosynthesis in both genotypes. Alpha‐tocopherol showed an important protective role against drought stress in Yousof cultivar as a relatively drought‐tolerant cultivar.     

Drought Stress in Grain Legumes during Reproduction and Grain Filling

Water scarcity is a major constraint limiting grain legume production particularly in the arid and semi‐arid tropics. Different climate models have predicted changes in rainfall distribution and frequent drought spells for the future. Although drought impedes the productivity of grain legumes at all growth stages, its occurrence during reproductive and grain development stages (terminal drought) is more critical and usually results in significant loss in grain yield. However, the extent of yield loss depends on the duration and intensity of the stress. A reduction in the rate of net photosynthesis, and poor grain set and grain development are the principal reasons for terminal drought‐induced loss in grain yield. Insight into the impact and resistance mechanism of terminal drought is required for effective crop improvement programmes aiming to improve resistance to terminal drought in grain legumes. In this article, the impact of terminal drought on leaf development and senescence, light harvesting and carbon fixation, and grain development and grain composition is discussed. The mechanisms of resistance, management options, and innovative breeding and functional genomics strategies to improve resistance to terminal drought in grain legumes are also discussed.     

土壤干旱对小麦绿色器官光合电子传递和光合磷酸化活力的影响

比较研究了开花后干旱胁迫下小麦(Triticum aestivum L.)不同绿色器官光合活性的变化，发现叶片光合对干旱胁迫的敏感性大于穗器官。在干旱胁迫下各器官叶绿体Hill反应活力、光合电子传递活性及光合磷酸化水平均下降，但下降幅度以旗叶叶片明显大于穗、穗下节间和叶鞘。暗示穗等非叶器官的光合作用具有较强的耐逆性。     

Temperature Effects on the Duration of the Spikelet Growth Phase and Spikelet Abortion in Barley

Above optimal temperatures in cereals, when measured in thermal time units (°Cd) can lead to underprediction of developmental events by models, particularly when cereals are sown late in Mediterranean-type environments or grown under subtropical conditions. Our objective was to assess the effects of high temperatures on the length of the spikelet growth phase, the number of spikelet nodes per spike and level of spikelet abortion in spring barley (Hordeum vulgare L.). The spikelet growth phase (from the end of the spikelet initiation phase to anthesis) of three cultivars, 'Bandulla', 'Schooner' and 'Weeah' was subjected to two temperature regimes. In the first of two sowings, the mean daily maximum temperature for one regime was 24 °C (maintained for 6 h daily) and the mean daily minimum was 8 °C (24/8 °C). The second temperature regime was 27/17 °C. Corresponding temperatures for the second sowing, which was subjected to slightly longer photoperiods, were 26/8 °C and 26/17°C, respectively. The duration of the spikelet growth phase was longer for 27/17 °C than for 24/8 °C in the first sowing when measured in calendar time (d). In the second sowing, the duration of the spikelet growth phase was slightly shorter under the higher temperature regime (d). The duration of the spikelet growth phase (in°Cd) was greatly increased by the higher temperature treatment in both sowings. In the first sowing, the percentage of aborted spikelets was greater at HT than at LT for Bandulla and Weeah at both sowing times and the duration of the spikelet growth phase increased with higher night temperatures, suggesting that length of the spikelet growth phase was not the sole factor responsible for the proportion of aborted spikelets.     

Drought Priming at Vegetative Growth Stage Enhances Nitrogen‐Use Efficiency Under Post‐Anthesis Drought and Heat Stress in Wheat

To study the effects of early drought priming at 5th‐leaf stage on grain yield and nitrogen‐use efficiency in wheat (Triticum aestivum L.) under post‐anthesis drought and heat stress, wheat plants were first exposed to moderate drought stress (drought priming; that is, the leaf water potential reached ca. ?0.9 MP a) at the 5th‐leaf stage for 11 days, and leaf water relations and gas exchange rates, grain yield and yield components, and agronomic nitrogen‐use efficiency (ANUE ) of the primed and non‐primed plants under post‐anthesis drought and heat stress were investigated. Compared with the non‐primed plants, the drought‐primed plants possessed higher leaf water potential and chlorophyll content, and consequently a higher photosynthetic rate during post‐anthesis drought and heat stress. Drought priming also resulted in higher grain yield and ANUE in wheat under post‐anthesis drought and heat stress. Drought priming at vegetative stage improves carbon assimilation and ANUE under post‐anthesis drought and heat stress and their combination in wheat, which might be used as a field management tool to enhance stress tolerance of wheat crops to multiple abiotic stresses in a future drier and warmer climate.     

Characterization of Barley Leaf Tolerance to Drought Stress by Chlorophyll Fluorescence and Electron Paramagnetic Resonance Studies

Two kinds of barley genotypes with various water‐stress tolerances, tolerant Cam/B1 and sensitive Maresi, were subjected to 10‐day soil‐drought stress in seedling and flag leaf developmental phases. After this time, both genotypes regardless of the growth stage showed a decrease in quantum yield of PSII photochemistry (Φ_PSII) upon stress treatment; however, this effect was stronger in the sensitive plants than in the tolerant ones. The drought stress in the flag leaf stage was associated with an increase in superoxide dismutase (SOD) level in both genotypes, whereas in seedlings, this effect was observed only for Maresi. The activity of other enzymes (catalase and peroxidase) was changed only in small degree. An increase in proline levels and activities of Δ¹‐pyrroline‐5‐carboxylate synthetase (P5CS) and ornithine delta‐aminotransferase (OAT) were observed independently of genotype and the phase of plant development, whereas the activity pyruvate dehydrogenase (PDH) decreased in tolerant genotype. Moreover, changes in the concentration of monocarbohydrates (glucose and fructose) and dicarbohydrates (saccharose, raffinose and maltose) were found: in seedlings, the amount of all soluble sugars increased, while in flag leaves decreased. The drought treatment resulted in a drop in starch level in the tolerant genotype, but in the sensitive one, the content of this substance increased in both developmental stages. EPR studies allowed the determination of the amount and character of organic radicals present in leaves. In control conditions, the content of these radical species was higher in the sensitive genotype than in tolerant one and decreased upon water stress, with the exception of flag leaves of the sensitive plant. Simulation procedure revealed four types of signals in the EPR spectra. One of them was attributed to a chlorophyll a cation and decreased upon drought. The second, ascribed to semiquinone radicals, reflected the redox balance disturbed by water deficit. The two remaining signals were connected with carbon‐centred radicals situated in the carbohydrate matrix. Their number was correlated with starch concentration.     

垄作栽培对啤酒大麦产量和品质的影响

探讨在等量灌水条件下垄作栽培对啤酒大麦产量和品质的影响。结果表明：（1）垄作栽培生育期长于传统平作栽培,单株穗数低于传统平作栽培,经济系数高于传统平作栽培,株高、穗粒数、产量影响趋势不一致,品种之间存在差异。（2）原麦品质：垄作栽培与传统平作栽培籽粒千粒重之间差异不显著,蛋白质含量之间差异显著,粒筛选率之间差异极显著水平,千粒重、蛋白质含量、筛选率均降低。（3）麦芽品质：垄作栽培麦芽蛋白质含量均较传统平作栽培降低,麦芽细粉浸出物、β-葡聚糖、a-氨基氮、a-淀粉酶、糖化力、可溶性氮之间差异不显著,库值含量之间差异达显著水平,但影响趋势不一致,品种之间存在差异。     

Fungal Endophytes Enhance Agronomically Important Traits in Severely Drought‐Stressed Barley

Crops often face severe and damaging local drought events, and in some regions, these episodes are predicted to become more frequent due to climate change. Some micro‐organisms have been shown to improve drought tolerance and improve yield in crop plants. Here, we show that fungal root endophytes isolated from a wild barley species (Hordeum murinum subsp. murinum) induced significant improvements in agronomic traits for a severely drought‐stressed barley cultivar grown in a controlled environment, including number of tillers, grain yield and shoot biomass. Five endophyte strains were tested, and the trait that showed the greatest significant difference in the drought‐stressed plants was the number of tillers, where all of the endophyte treatments induced a greater number of tillers per plant. However, except in one case, the mean dry root weight for all plants was greater in the control plants, indicating preferential allocation of resources to aboveground parts in the endophyte treatments. Results were not consistent across all endophyte treatments, with some endophytes performing much better than others. As these growth studies were conducted using soil‐based compost, the results may translate to the field and suggest that some of these endophytes have potential as barley inoculants in arid growing conditions.     

Grain Weight Reductions in Wheat Associated with Semidwarfism: an Analysis of Grain Weight at Different Positions Within the Spike of Near-isogenic Lines

Semidwarf genes have made a major contribution to grain yield gains in wheat during this century. These genes have frequently been reported to increase grain number but decrease grain size. To analyse possible causes of this negative association between number of grains per head and mean grain weight, grains from different positions within the spikes were individually weighed in three isolines (standard-height, SH; semidwarf, SD; and dwarf, DD) of the spring wheat cultivar Maringa grown under field conditions. Dwarfing genes affected both grain number and grain weight. Increasing doses of Rht genes resulted in a decreased average grain weight (ca. 22 %) but in a greater increase in number of grains per m² (ca. 40%). Since the isogenic lines did not differ in either the number of spikes per m² or number of spikelets per spike, the frequencies of appearance of grains in the distal positions (florets 3 and 4) of the central spikelets and in the top spikelets of the spike was much higher for the SD and DD lines than for the SH line. Averaging across spikelets, basal grains (grains 1 and 2) of SH, SD and DD lines represented ca. 82, 73, and 69 %, respectively, of the total number of grains. On the other hand, grain 4 was almost absent in the SH line but represented around 10 % of the grains in the DD line. Averaging across lines and spikelets, basal grains weighed ca. 30 mg, which is approximately 20 % heavier than grain 3 (ca. 25 mg) and 85 % heavier than grain 4 (ca. 16 mg). Although there was a clear trend for a greater grain weight in the SH line (with no major differences between the SD and DD line) when basal grains were considered, grains 3 and 4 did not show any clear difference between lines. The fact that genotypes with Rht alleles had grains in the distal positions with a much higher frequency of appearance than in the SH line could be the main reason for their lower average grain weight. Therefore, the negative relationship between grain number and average grain weight might be not reflecting that competition for assimilates among grains of the DD and SD lines is stronger than that in the SH line.     

Yield Responsiveness in Two- and Six-Rowed Barley Grown in Contrasting Nitrogen Environments

Two‐ and six‐rowed barley with different intrinsic ability to produce tillers and kernels per ear, would differ in responsiveness to nitrogen availability with environmental improvements. Two field experiments were carried out to elucidate how nitrogen supply (N₄₀ and N₁₅₀) affects yield and its components in two‐ and six‐rowed barley. High nitrogen increased aboveground dry matter at anthesis, by improving cumulative solar radiation intercepted by the crop, determining an increased dry‐matter production at maturity without changes in harvest index. In both barley types, variations in grain yield were explained by changes in kernels per unit land area rather than by differences in the average kernel weight. However, changes in the number of kernels were due to variations in the number of ears per m² in two‐rowed barley and the number of kernels per ear in six‐rowed barley. Ears per unit area showed a greater responsiveness in two‐ than in six‐rowed barley due to a higher nitrogen supply treatment, associated with their intrinsic higher tillering capacity, while the number of kernels per ear was more responsive in six‐ than in two‐rowed types. The fact that responses to nitrogen by the number of kernels per unit land area in two‐ and six‐rowed barley is better explained by different yield sub‐components, allows the speculation that the critical period for yield determination would differ between barley types.     

Evaluation of Drought-Related Traits and Screening Methods at Different Developmental Stages in Spring Barley

Despite intensive research and breeding efforts, the physiological and quantitative genetic bases of drought tolerance are still poorly understood. The comparison of results obtained from different sources is also complex, because different testing methods may lead to controversial conclusions. This report discusses various drought stress experiments (hydroponics and in soil) in which the plant tolerance was studied at different developmental stages. Tests were performed in the germination, seedling and adult plant stages on the parental lines of five well-known barley-mapping populations. The results suggest that drought tolerance is a stage-specific trait and changes during the life cycle. The effect of drought stress depended not only on the duration and intensity of water deficiency, but also on the developmental phase in which it began. To induce the same type of stress and to obtain comparable tolerance information from the replications, it is recommended that drought stress should be induced at the same growth stage. Correlations between the traits, commonly associated with improved drought resistance (high relative water content under stress, proline accumulation, osmoregulation) with stress tolerance indexes, are also presented, while the advantages and disadvantages of the most frequently used screening methods are discussed.     

Independent and Combined Effects of Soil Warming and Drought Stress During Anthesis on Seed Set and Grain Yield in Two Spring Wheat Varieties

Increase in soil temperature together with decrease in soil moisture during anthesis of spring wheat (Triticum aestivum L) crops is predicted to occur more frequently in a future climate in Denmark. The objective of this study was to investigate the responses of two Danish spring wheat varieties (Trappe and Alora) to soil warming (H), drought (D) and both (HD) during anthesis. The plants were grown in pots in a climate‐controlled glasshouse. In H, the soil temperature was increased by 3 °C compared with the control (C). In both D and HD treatments, the plants were drought‐stressed by withholding irrigation until all of the transpirable soil water had been depleted in the pots. Results showed that, particularly under D treatment, Alora depleted soil water faster than Trappe. In both varieties, flag leaf relative water content (RWC) was significantly lowered, while spikelet abscisic acid (ABA) concentration was significantly increased by D and HD treatments. Compared with the C plants, D and HD treatments significantly reduced ear number, ear to tiller ratio, shoot biomass, grain yield, harvest index and seed set but hardly affected tiller number and 1000‐kernel weight, whereas H treatment alone only decreased shoot biomass and reduced seed set. When analysed across the varieties and the treatments, it was found that the reduction in seed set was closely correlated with the increase in spikelet ABA concentration, indicating that D and HD treatments induced greater spikelet ABA concentrations might have caused seed abortion. It was concluded that the grain yield reduction under D and HD treatments during anthesis in spring wheat is ascribed mainly to a lowered seed set and wheat varieties (i.e. Alora) with more dramatic increase in spikelet ABA concentration are more susceptible to D and HD treatment.