Short-term responses of leaf growth rate to water deficit scale up to whole-plant and crop levels: an integrated modelling approach in maize

Physiological and genetic studies of leaf growth often focus on short-term responses, leaving a gap to whole-plant models that predict biomass accumulation, transpiration and yield at crop scale. To bridge this gap, we developed a model that combines an existing model of leaf 6 expansion in response to short-term environmental variations with a model coordinating the development of all leaves of a plant. The latter was based on: (1) rates of leaf initiation, appearance and end of elongation measured in field experiments; and (2) the hypothesis of an independence of the growth between leaves. The resulting whole-plant leaf model was integrated into the generic crop model APSIM which provided dynamic feedback of environmental conditions to the leaf model and allowed simulation of crop growth at canopy level. The model was tested in 12 field situations with contrasting temperature, evaporative demand and soil water status. In observed and simulated data, high evaporative demand reduced leaf area at the whole-plant level, and short water deficits affected only leaves developing during the stress, either visible or still hidden in the whorl. The model adequately simulated whole-plant profiles of leaf area with a single set of parameters that applied to the same hybrid in all experiments. It was also suitable to predict biomass accumulation and yield of a similar hybrid grown in different conditions. This model extends to field conditions existing knowledge of the environmental controls of leaf elongation, and can be used to simulate how their genetic controls flow through to yield.     

Ammonium, calcium, and leaf senescence in rice

The possible involvement of calcium in the regulation of ammonium-promoted senescence of detached rice leaves was investigated. Calcium effectively reduced ammonium-promoted senescence of detached rice leaves. The effect of ammonium on the senescence was also significantly reduced by the calcium ionophore A23187. Ammonium-promoted senescence of detached rice leaves may be mediated through blocking the entrance of calcium ions into the cytosol.     

Leaf area and crop growth in relation to phenology of barley in Mediterranean environments

In a barley/livestock farming system of northern Syria, high biomass production in addition to high grain yield is desirable. The aim of this study was to assess the effect of environment and phenology on growth and yield of barley in northern Syria. Leaf area duration (LAD), specific leaf area (SLA), crop growth rate (CGR) on a thermal time basis, and biological and grain yields were compared for entries representing three contrasting development patterns: early heading spring types (pattern A), medium early heading winter types (pattern B), and late heading spring types (pattern C). The experiment was conducted in 1988/89 at two sites: Tel Hadya (TH, 250 mm precipitation) and Breda (BR, 180 mm). Cold damage occurred in winter and, especially at BR, drought stress developed in spring.At the two sites, development was similar, but yields at TH were twice those at BR. This was related to a longer LAD and a faster CGR in spring. Development pattern affected growth. A long vegetative phase (pattern B) resulted in small leaves with a low SLA in winter, probably due to a slow leaf extension rate. Since cold tolerance and profuse tillering compensated for the small leaf size, pattern B had on average a longer LAD than pattern A. Pattern C had a longer LAD than pattern A because of a longer crop duration. This long duration had a negative effect on yield, so LAD was poorly related to yield. Development in spring was associated with CGR. Pattern C had a slow CGR and low yields; pattern B had the fastest CGR, but the yield advantage over pattern A was not significant. These results suggest that early heading winter barley, which combines long LAD with fast spring CGR, may give the best performance in a barley/livestock farming system in northern Syria.     

大麦叶面积指数模拟模型

准确模拟叶面积指数是作物生长模拟模型预测作物生长和产量的关键.本文通过系统分析扬州和武汉地区不同大麦品种高产群体叶面积指数变化动态,建立了大麦群体的叶面积指数模拟模型.大麦叶面积指数是品种叶面积指数扩展的遗传参数和气温日较差、日照时数、辐射量等气候因子及水肥丰缺因子的函数.孕穗抽穗期最大叶面积指数与该期最适叶面积指数是不同的概念,二者之间存在着极显著差异.利用扬州、南京和昆明地区不同品种的播期试验及氮肥试验资料对模型进行了检验,结果表明,模型对大麦叶面积指数的模拟效果较好,模拟值与观测值吻合度高,根均方差RMSE介于0.742～2.865,平均值为1.348.对模拟值与观测值进行y=x的线性回归分析,相关系数R²介于0.511～0.954,均呈极显著正相关.     

基于作物生长监测诊断仪的双季稻叶片氮含量和氮积累量监测

为了验证作物生长监测诊断仪(CGMD)监测双季稻氮素营养指标的准确性和适用性,构建基于CGMD的双季稻叶片氮含量(LNC)和氮积累量(LNA)的监测模型。选用8个不同早、晚稻品种,设置4个不同施氮水平,利用CGMD采集冠层差值植被指数(DVI)、归一化植被指数(NDVI)和比值植被指数(RVI),同步利用ASD FH2高光谱仪采集冠层光谱反射率,并计算DVI、NDVI和RVI;通过比较CGMD和ASD FH2采集的冠层植被指数变化特征,验证CGMD的测量精度,构建基于CGMD的LNC和LNA监测模型,并利用独立试验数据对模型进行检验。结果表明: 早、晚稻LNC、LNA、DVI、NDVI和RVI随施氮水平的增加而增大,随生育进程的推进呈先升后降的趋势;CGMD与ASD FH2采集的DVI、NDVI和RVI间拟合的决定系数(R²)分别为0.9350、0.9436和0.9433,表明CGMD的测量精度较高,可替代ASD FH2采集冠层植被指数。基于CGMD的3个冠层植被指数相比,NDVI_CGMD与LNC的相关性最高,RVI_CGMD与LNA的相关性最高;基于NDVI_CGMD的指数模型可较准确地预测LNC,模型R²为0.8581～0.9318,模型检验的均方根误差(RMSE)、相对均方根误差(RRMSE)和相关系数(r)分别为0.1%～0.2%、4.0%～8.5%和0.9041～0.9854;基于RVI_CGMD的幂函数模型可较准确地预测LNA,模型R²为0.8684～0.9577,模型检验的RMSE、RRMSE和r分别为0.37～0.89 g·m^-2、6.7%～20.4%和0.9191～0.9851。与化学分析方法相比,利用CGMD可便捷准确地获取早、晚稻的LNC和LNA,在双季稻丰产高效栽培和氮肥精确管理中具有应用价值。     

Dynamic model of the response of a vegetative grass crop to light, temperature and nitrogen

Abstract A previously described growth model of the vegetative grass crop is extended to include a simple representation of the root system, uptake of nitrogen from a soil nitrogen pool, and response to fertilizer application. The model simulates the processes of light interception, photosynthesis, partitioning of new growth, leaf area expansion, growth and maintenance respiration, ageing of plant tissues, senescence, recycling of substrates from senescing tissues, nitrogen uptake by the plant, leaching, mineralization, and fertilizer application. A principal component of the model, nitrogen uptake, is assumed to depend positively on plant carbon substrate concentration and soil nitrogen concentration, and to be inhibited by plant nitrogen substrate concentration. The dynamic responses to different levels of soil nitrogen, of shoot and root growth, nitrogen uptake and root activity, carbon and nitrogen plant substrate concentrations, and the fraction of substrate carbon used by the shoots, are examined; realistic behaviour is observed. The model predicts nitrogen fertilizer responses of yield and plant nitrogen content, which are compared directly with experimental data; good agreement is obtained.     

Directed overexpression of PHYA locally suppresses stem elongation and leaf senescence responses to far-red radiation

Ectopic overexpression of an oat PHYA cDNA in tobacco under the cauliflower mosaic virus 35S promoter results in plants with reduced morphological responses to far-red radiation (FR). We have tested the hypothesis that it is possible to molecularly ‘mask’ steins and leaves to FR-induced elongation and senescence responses by targeting the over-expression of PHYA with appropriate promoters. Oat PHYA was expressed in tobacco (Nicotiana tabacum L. cv Xanthi) under the 35S and two Arabidopsis promoters: UBQ1 and CAB. The internodes of wild type, UBQ:PHYA, and CAB:PHYA plants, which exhibited little or no oat PHYA overexpression, responded to localized FR treatments with a marked increase in elongation. In contrast, 35S:PHYA plants, which overexpressed PHYA to high levels in all parts of the shoot, did not respond to FR treatments directed to their stems. Leaf senescence responses to FR were remarkably localized, and sensitivity to FR was also inversely correlated with the local PHYA expression level. Thus, chlorophyll content, specific leaf weight, and nitrate reductase activity in leaf spots treated with FR were highly reduced in wild type and UBQ:PHYA plants, but not in the CAB:PHYA and 35S:PHYA counterparts. Our results suggest that it may be feasible to obtain transgenic crop plants in which certain organs or tissues are made ‘blind’ to phytochrome-perceived signals of canopy density, but whose general photomorphogenic competence is not greatly disturbed.     

Water stress, ammonium, and leaf senescence in detached rice leaves

Ammonium accumulation in relation to water stress-promoted senescence of detached rice leaves was investigated. The effect of water stress on the senescence of detached rice leaves is associated with the accumulation of ammonium. The accumulation of ammonium in detached rice leaves by water stress is attributed to a decrease in glutamine synthetase activity. Ammonium accumulation in detached rice leaves, induced by water stress, was accompanied by an increase in tissue sensitivity to ethylene which, in turn, accelerated leaf senescence.     

Plant development controls leaf area expansion in alfalfa plants competing for light

Background and Aims

The growth of crops in a mixture is more variable and difficult to predict than that in pure stands. Light partitioning and crop leaf area expansion play prominent roles in explaining this variability. However, in many crops commonly grown in mixtures, including the forage species alfalfa, the sensitivity and relative importance of the physiological responses involved in the light modulation of leaf area expansion are still to be established. This study was designed to assess the relative sensitivity of primary shoot development, branching and individual leaf expansion in alfalfa in response to light availability.

Methods

Two experiments were carried out. The first studied isolated plants to assess the potential development of different shoot types and growth periods. The second consisted of manipulating the intensity of competition for light using a range of canopies in pure and mixed stands at two densities so as to evaluate the relative effects on shoot development, leaf growth, and plant and shoot demography.

Key Results

Shoot development in the absence of light competition was deterministic (constant phyllochrons of 32·5 °Cd and 48·2 °Cd for primary axes and branches, branching probability of 1, constant delay of 1·75 phyllochron before axillary bud burst) and identical irrespective of shoot type and growth/regrowth periods. During light competition experiments, changes in plant development explained most of the plant leaf area variations, with average leaf size contributing to a lesser extent. Branch development and the number of shoots per plant were the leaf area components most affected by light availability. Primary axis development and plant demography were only affected in situations of severe light competition.

Conclusions

Plant leaf area components differed with regard to their sensitivity to light competition. The potential shoot development model presented in this study could serve as a framework to integrate light responses in alfalfa crop models.     

Modeling leaf senescence of deciduous tree species in Europe

Autumnal leaf senescence signals the end of photosynthetic activities in temperate deciduous trees and consequently exerts a strong control on various ecological processes. Predicting leaf senescence dates (LSD) with high accuracy is thus a prerequisite for better understanding the climate–ecosystem interactions. However, modeling LSD at large spatial and temporal scales is challenging. In this study, first, we used 19972 site‐year records (848 sites and four deciduous tree species) from the PAN European Phenology network to calibrate and evaluate six leaf senescence models during the period 1980–2013. Second, we extended the spatial analysis by repeating the procedure across Europe using satellite‐derived end of growing season and a forest map. Overall, we found that models that considered photoperiod and temperature interactions outperformed models using simple temperature or photoperiod thresholds for Betula pendula, Fagus sylvatica and Quercus robur. On the contrary, no model displayed reasonable predictions for Aesculus hippocastanum. This inter‐model comparison indicates that, contrary to expectation, photoperiod does not significantly modulate the accumulation of cooling degree days (CDD). On the other hand, considering the carryover effect of leaf unfolding date could promote the models’ predictability. The CDD models generally matched the observed LSD at species level and its interannual variation, but were limited in explaining the inter‐site variations, indicating that other environmental cues need to be considered in future model development. The discrepancies remaining between model simulations and observations highlight the need of manipulation studies to elucidate the mechanisms behind the leaf senescence process and to make current models more realistic.     

Use of lysophosphatidylethanolamine, a natural lipid, to retard tomato leaf and fruit senescence

We studied the influence of lysophosphatidylethanolamine (LPE) on the pattern and rate of ethylene production and respiration of tomato ( Lycopersicon esculentum cv. H7155) leaflets and fruit. Leaflets that had been senescing on the plant showed a climacteric-like rise in ethylene production but not in respiration rate which decreased continuously with leaf age. Detached leaflets had a climacteric-like pattern in respiration whether they were incubated in complete darkness or in light. Detached leaflets incubated in the dark had higher rates of ethylene production and CO₂ evolution than did light-incubated leaves. There was no change in the pattern of ethylene production or CO₂ evolution as a result of LPE treatment. However, LPE-treated attached and detached leaflets had consistently lower rates of CO₂ evolution. The reduction in CO₂ evolution by LPE was most pronounced at the climacteric-like peak of the detached leaves. LPE-treated leaflets had a higher chlorophyll content and fresh weight and lower electrolyte leakage than the control. LPE-treated fruits had lower rates of ethylene and CO₂ production than did the control. LPE-treated fruits also had higher pericarp firmness and lower electrolyte leakage than the control. The results of the present study provide evidence that LPE is able to retard senescence of attached leaves and detached leaves and tomato fruits. Several recent studies suggest that lysolipids can act in a specific manner as metabolic regulators. Our results suggest a specific role of lysolipid LPE in aging and senescence     

基于叶干质量比的杉木比叶面积估算模型的构建

随着叶片功能性状研究的不断深入, 通过简单易测量的叶片指标, 同时探究植物生活史权衡对策和估算林分生产力的研究需求日益增长, 例如叶干质量比(LDMC)和比叶面积(SLA)的相互转换。杉木(Cunninghamia lanceolata)是亚热带重要的常绿针叶树种, 基于LDMC对杉木SLA进行估算, 能够为核算SLA提供途径, 为机理解释和生产估算构建连接途径, 为小区域到大尺度、精算到估算搭建桥梁。该研究在湖南会同和河南信阳两个杉木生长区, 对处于不同小生境(坡向、坡位和冠层深度)以及不同生活史(林龄和叶龄)的叶片进行抽样和采集, 通过测得不同叶龄的单叶LDMC和SLA, 初步探究在不同因子下两个性状值的分布差异, 进一步基于LDMC构建SLA估算模型并讨论以叶龄为差分因子对模型的影响。结果表明: 1)杉木SLA平均值为(103.15 ± 69.54) cm ²·g ^-1, LDMC为0.39 ± 0.11; 2)杉木LDMC和SLA可用非线性模型进行估算, 模型符合估算要求; 3)其中一年生叶的拟合效果最好, 老叶(大于二年生叶)的拟合优度较低, 老叶较低的SLA (52.28-75.74 cm ²·g ^-1)可能暗示LDMC的变化保持相对独立性。该研究基于杉木LDMC的SLA估算模型可信且有效, 且不同叶龄对LDMC和SLA的影响可能预示着杉木叶片的响应敏感性和生活史权衡策略。     

Cytokinin biochemistry in relation to leaf senescence. VIII. Translocation, metabolism and biosynthesis of cytokinins in relation to sequential leaf senescence of tobacco

Cytokinin bases (zeatin and dihydrozeatin) and ribosides (zeatin riboside and dihydrozeatin riboside) were identified as major cytokinins in tobacco xylem sap by radioimmunoassay. When ³H-labelled zeatin riboside or dihydrozeatin riboside were supplied to tobacco plants via the xylem, leaves of differing maturity did not differ appreciably in level of radioactivity or in metabolism of the cytokinin. The major metabolites of zeatin riboside in leaves were adenine, adenosine and adenine nucleotides, whereas that of dihydrozeatin riboside was dihydrozeatin 7-glucoside. Incorporation of [¹⁴C]adenine into zeatin was evident in upper green leaves. indicating that young leaves have the capacity to synthesize cytokinins in situ. In contrast, fully expanded green leaves and senescing tobacco leaves exhibited little or no incorporation of [¹⁴C]adenine into cytokinins. This difference in cytokinin biosynthetic capacity may contribute to the differing cytokinin levels in leaves of different matirity, and may participate in control of sequential leaf senescence in tobacco.     

胡杨枝芽生长特征及其展叶物候特征

以5个不同发育阶段的胡杨(Populus euphratica Oliv.)个体为研究对象,观测记录了枝芽展叶物候、枝芽生长特征和叶形变化的空间分布规律。结果表明:不同发育阶段的胡杨个体以及同一个体树冠的不同层次,其枝芽生长及其展叶物候期表现出不同的时空特征。随着树龄的增加和树冠层次的增高(由基向顶),当年新生枝条长度、枝条叶片数和叶形指数逐渐减小,但叶面积和叶片干重逐渐增大。5个不同发育阶段胡杨个体均表现出展叶物候始于树冠顶层,依次向下结束于树冠基部;展叶物候期共性表现在枝芽萌动期均在4月上旬,起始展叶期集中在4月中旬,展叶终期则在5月上旬到下旬;树龄较大的个体其枝芽萌动期、起始展叶期、展叶终期较树龄较小的个体早;其枝芽萌动期到展叶终期的时间进程较树龄较小的个体短;不同发育阶段的个体枝芽萌动期出现的时间较为离散,起始展叶期和展叶终期出现的时间较为集中。相关分析表明,出叶周期与枝条长度、枝条叶片数量和叶形指数呈极显著正相关,与叶面积和叶片干重呈显著负相关。     

Cytokinin biochemistry in relation to leaf senescence. VI. Effect of nitrogenous nutrients on cytokinin levels and senescence of tobacco leaves

The effect of nitrogenous nutrients on endogeneous cytokinins and senescence of tobacco leaves was investigated. Ammonium nitrate was the most effective in retarding senescence and its activity was attributed principally to NH₄⁺ ions. Repeated applications or a continuous supply of ammonium nitrate was required for maximal retardation of tobacco leaf senescence. Ammonium nitrate solution supplied via the petioles reduced the senescence retarding effect of dihydrozeatin applied directly to the laminae of detached tobacco leaves. Ammonium nitrate also elevated the endogenous levels of cytokinins (especially zeatin and dihydrozeatin) particularly in growing tobacco leaves excised from near the apex of the plant. Ammonium nitrate induced retardation of leaf senescence may be mediated at least partly by its effect on foliar cytokinin content.     

基于WOFOST作物生长模型的冬小麦干旱影响评估技术

为了反映作物与干旱的相互关系,人为再现干旱灾害对作物产量的影响程度,选择华北地区冬小麦干旱灾害为研究对象,对作物生长模型WOFOST在区域上进行适应性进行分析、检验的基础上,然后利用区域作物模型实现干旱灾害对作物影响定量分析和动态评估。以减产率和气象条件作为灾害严重程度划分的标准,利用数值模拟试验,确定导致减产的主要气象因子及其量值,对研究区干旱灾害进行影响评估,包括典型灾害年份影响评估和年代际灾害影响评估,并给出了评估结果。     

杉木叶形态特征与叶面积估算模型

植物叶片是碳水交换和能量平衡过程最重要的场所,是农林生产经营中的模型估算以及物种结构变异-功能适应机制分析的关键参考量。采用游标卡尺和手持叶面积仪,测量杉木(Cunninghamia lanceolata)单叶叶长(LL)、最大叶宽(LW_(max))、最大叶厚(LT_(max))3个直测指标,和叶面积(LA)、平均叶宽(LW_(mean))、平均叶厚(LT_(mean))、叶延长率(LE)和叶周长(LP)5个间接转算指标。分析8个形态学指标的统计分布及其相关性,用多变量线性回归模型和非线性回归指数模型对7个形态学指标和杉木单叶叶面积进行拟合,结果表明:杉木单叶面积大部分值(95%CI)分布在0.758—0.836 cm~2,其叶面积的变异程度最大(CV=0.513),叶长、叶宽与叶面积相关性达到极显著(r=0.896,0.682)。拟合LA的多元线性模型为:Y=-0.388+0.165X_1-0.023X_2+1.453X_3(R~2=0.981,SE=0.053),X_1—X_3分别为LP、LE、LW_(mean)。从简便性上考虑,LL的单变量指数模型适合对LA进行估算:LA=0.1×(1+LL)~(1.398)(R~2=0.77,!~2=0.39)。研究结果为准确估测其他叶片功能性状指标提供了方法,为杉木叶面积估算模型提供了基础数据。