夏玉米叶片气体交换参数对干旱过程的响应

目前已经开展了大量的干旱对作物叶片气体交换参数影响的研究,但关于作物叶片气体交换参数对干旱过程的响应及其关键阈值的研究仍较少。基于夏玉米七叶期开始的5个初始水分梯度的长时间持续干旱模拟实验资料,分析了不同强度持续干旱过程中夏玉米叶片气体交换参数(净光合速率Pn,气孔导度Gs,蒸腾速率Tr,胞间CO_2浓度Ci和气孔限制值Ls)的变化规律及其关键阈值。结果表明,玉米的净光合速率(Pn),蒸腾速率(Tr)和气孔导度(Gs)在干旱发生初期呈大幅度下降,但随着干旱持续会出现一定的适应性。利用统计容忍限方法确定了夏玉米拔节期Pn,Tr和Gs响应干旱的临界土壤相对湿度(0—30cm)分别为53%,51%和48%,对应的临界叶含水率分别为81.8%,81.3%和81.2%。夏玉米光合作用由气孔限制向非气孔限制转换的0—30cm土壤相对湿度均为44%±2%,对应的叶含水率均为77.6%±0.3%。研究结果可为夏玉米干旱发生发展过程的监测预警提供依据。     

华北夏玉米干物质分配系数对干旱胁迫的响应

干物质分配系数反映作物各器官干物质的分配与积累,研究干物质分配系数对干旱胁迫的响应,是研究干旱胁迫对作物生长发育影响的基础.本文基于华北夏玉米主产省山东、河北和山西3个试验点2013—2015年田间水分控制试验资料,建立了夏玉米苗期、抽雄期、灌浆期3个主要发育阶段叶、茎、穗的干物质分配系数与土壤相对湿度的定量关系模型,分析了叶、茎、穗干物质分配系数对不同程度干旱胁迫的响应.结果表明: 3个阶段叶、茎、穗的干物质分配系数与土壤相对湿度均呈显著的一元二次关系.干旱胁迫下,叶片向外转运的干物质相对减少,叶干物质分配比例增加,并且在轻、中度干旱胁迫时的灌浆期(叶干物质分配系数增加0.04~0.09)以及重度干旱胁迫时的抽雄期(叶干物质分配系数增加0.17)响应最敏感.穗干物质分配系数对干旱胁迫表现为负响应,干旱胁迫越严重,分配系数越小,轻-重度干旱胁迫使穗干物质分配系数减小0.08~0.34.茎干物质分配系数对干旱胁迫的响应总体表现为灌浆期(正响应)＞抽雄期(负响应)＞苗期(负响应).     

施肥对干旱胁迫下夏玉米根系提水的调节作用研究

以耐旱性玉米品种'郑单958号'为材料,采用两室分根土培装置,通过时域反射计(TDR)对上下土层土壤含水量进行控制和观测,研究施肥对干旱胁迫条件下玉米根系提水作用的影响.结果表明,玉米根系在土层上干下湿条件下(即上下层土壤存在一定水势差时)存在明显提水作用;玉米根系提水量在整个生育期呈单峰变化,并以吐丝期最大;上层土壤施肥可以调节玉米根系提水作用强弱,整个生育期根系总提水量表现为NP配施>单施P>CK>单施N, NP配施处理全生育期单株提水量(1 948.6 g)分别是单施P处理、CK和单施N处理的1.5倍、3.1倍和3.5倍.玉米整个生育期根系总提水量与收获期不同层次根系干重和体积存在极显著正相关关系,也与其地上部分生物量和籽粒产量呈极显著或显著正相关关系.可见,玉米根系的提水作用强弱因生育期和施肥处理而变化,施肥主要通过影响根系生长来调节其提水作用;在一定水分环境条件下,玉米根系提水作用能促进作物生长,提高其籽粒产量.     

干旱胁迫对丹参叶片气体交换和叶绿素荧光参数的影响

研究了干旱处理15d后,大叶型丹参和小叶型丹参2个品种幼苗气体交换和叶绿素荧光参数的变化.结果表明:在干旱胁迫15d后,大叶型丹参叶片净光合速率(Pn)和PSⅡ最大光化学效率(Fv/Fm)分别下降了66.42%和10.98%,而小叶型丹参的Pn和Fv/Fm分别下降了29.32%和5.47%,干旱胁迫对大叶型丹参Pn和Fv/Fm的影响明显大于小叶型丹参.小叶型丹参Pn下降主要由气孔因素造成,而大叶型则主要由非气孔因素所致.干旱胁迫使丹参叶片的气孔导度(Gs)下降,但明显诱导了水分利用效率(WUE)、非光化学猝灭系数(qN)和光呼吸速率与净光合速率比率(Pr/Pn)的增加,以提高干旱胁迫抗性.其中小叶型丹参的增幅明显大于大叶型丹参.表明小叶型丹参的抗干旱胁迫能力更强.     

ABA对玉米响应干旱胁迫的调控机制

玉米(Zea mays L.)在生长期常常受到干旱的胁迫,而脱落酸(ABA)调节的生长响应是植物对逆境信号的一个基本反应. 本文对近年来国内外有关ABA提高玉米抗氧化防护系统,保护细胞免受氧化损伤,增加可溶性渗透剂(如脯氨酸)维持细胞内的水分,以及与其它激素相互作用影响玉米器官发育等的研究进展进行综述,以了解ABA调控玉米根系、叶片和籽粒的耐旱机理.     

干旱胁迫下氮素营养与根信号在气孔运动调控中的协同作用

干旱胁迫下根系与地上部分之间的信息传递可使植物叶片及时感知土壤水势变化,从而使植物在没有真正受到干旱伤害时即可做出主动、快速的抗旱应答反应,而在这一过程中,脱落酸（abscisic acid,ABA）和pH起着关键的作用。本研究表明。干旱胁迫下鸭趾草（Commelina communis L．）、番茄（Lycopersicon esculentum Mill．）和向日葵（Helianthus annuus L．）木质部汁液中pH的变化很不相同,且该pH变化和木质部汁液中硝态氮离子浓度的变化没有直接的关系;然而,饲喂实验表明,无论对于何种植物,蒸腾流中硝态氮离子浓度的增加都可有效地增加气孔对ABA的敏感度;分根实验进一步表明,土壤中硝态氮营养的增加可明显提高气孔对根信号的敏感度。以上结果说明,氮素营养可以和根信号相互作用共同操纵气孔运动。     

干旱胁迫下氮素营养与根信号在气孔运动调控中的协同作用

干旱胁迫下根系与地上部分之间的信息传递可使植物叶片及时感知土壤水势变化, 从而使植物在没有真正受到干旱伤害时即可做出主动、快速的抗旱应答反应, 而在这一过程中, 脱落酸(abscisic acid, ABA)和pH起着关键的作用。本研究表明, 干旱胁迫下鸭趾草(Commelina communis L.)、番茄(Lycopersicon esculentum Mill.)和向日葵(Helianthus annuus L.)木质部汁液中pH的变化很不相同, 且该pH变化和木质部汁液中硝态氮离子浓度的变化没有直接的关系; 然而, 饲喂实验表明, 无论对于何种植物, 蒸腾流中硝态氮离子浓度的增加都可有效地增加气孔对ABA的敏感度; 分根实验进一步表明, 土壤中硝态氮营养的增加可明显提高气孔对根信号的敏感度。以上结果说明, 氮素营养可以和根信号相互作用共同操纵气孔运动。     

干旱胁迫对玉米苗期叶片光合作用和保护酶的影响

以玉米品种郑单958(抗旱性强)和陕单902(抗旱性弱)为材料,采用盆栽控水试验,设置3个干旱处理(轻度干旱,中度干旱,重度干旱)和正常灌水,研究了干旱胁迫对玉米苗期叶片光合速率、叶绿素荧光以及相关生理指标的影响。结果表明:(1)干旱胁迫下2个品种叶片净光合速率(P_n)和气孔导度(G_s)显著下降,胞间CO₂浓度(C_i)出现了先下降后上升,而气孔限制值(L_s)上升后下降,说明中度干旱胁迫下叶片P_n下降是气孔因素引起的,重度干旱胁迫下P_n降低主要由非气孔因素引起的。(2)随着干旱胁迫的加剧,2个品种叶片光系统Ⅱ(PSⅡ)的实际量子产量(φPSⅡ)、电子传递速率(ETR)和光化学猝灭(qP)一直下降,而非光化学猝灭(qN)上升后下降,说明中度干旱下热耗散仍是植株重要光保护机制,重度干旱时叶片光合电子传递受阻,PSⅡ受到损伤。(3)干旱胁迫下2个品种叶片的超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)活性先升高后降低,而丙二醛(MDA)含量一直升高,说明干旱胁迫初期对保护系统酶活性升高有诱导作用,重度胁迫下活性氧清除酶的活性下降,导致细胞膜伤害。这些结果暗示,轻度和中度干旱胁迫下2个玉米品种通过减少光捕获、热耗散和酶活性调节协同作用稳定了光合机构功能,是P_n下降的气孔限制因素;而重度干旱胁迫下光系统Ⅱ和抗氧化酶系统损伤,是P_n下降的非气孔限制因素;郑单958的各生理参数比陕单902受旱影响小,干旱胁迫下仍具有较高的光合效率和较强的保护酶活性是郑单958抗旱的主要生理原因。     

试论麦类作物非水力根信号与生活史对策

从植物非水力根信号的生理调节作用和生活史进化解度看,在水分亏缺条件下,自然选择会导致植物产生大量根系以增加对水分的竞争能力,而浅根系则可在干旱来临时,以快速反应的根信号来调节和平衡植株水分状况,度过干旱时期。但是,自然选择压力下的植物特征往往不利于作物籽粒产量这一种群水平上的属性的改善。作物产量的提高过程是一个不断加强的人工选择过程。在作物生产中,作物水环境得到了改善,强大的多年生竞争者基本消失,     

施磷对干旱胁迫下箭竹根际土壤养分及微生物群落的影响

以箭竹及其根际土壤作为研究对象,采用两因素随机区组实验,设置2种水分处理（正常浇水和干旱胁迫）和2种施磷量处理（施磷和不施磷）,探究施磷对干旱胁迫下箭竹根际土壤养分及微生物群落结构和多样性的影响。结果表明：（1）干旱胁迫显著降低了箭竹根际土壤中微生物量碳、可溶性有机氮和有效磷的含量,虽对箭竹根际土壤微生物群落的多样性无显著影响,但显著降低了箭竹根际土壤中总PLFA（phospholipid fatty acid contents）的含量和真菌、细菌、革兰氏阳性菌与革兰氏阴性菌的PLFA含量以及革兰氏阳性菌/革兰氏阴性菌的PLFA比值,显著改变了箭竹根际土壤微生物群落结构,结果显著降低了箭竹的生物量。（2）施磷显著增加了受旱箭竹根际土壤中微生物量碳和有效磷的含量,虽大体上对受旱箭竹根际土壤微生物群落的多样性无显著影响,但显著增加了受旱箭竹根际土壤中总PLFA和真菌PLFA的含量,并在一定程度上增加了细菌、革兰氏阳性菌、革兰氏阴性菌和放线菌的PLFA含量以及革兰氏阳性菌/革兰氏阴性菌和真菌/细菌的PLFA比值,也在一定程度上改善了受旱箭竹根际土壤微生物群落结构,从而改善受旱箭竹的生长。（3）主成分分析表明,干旱对箭竹根际土壤微生物群落结构的影响显著,而施磷的影响不明显。（4）相关分析发现,箭竹根际土壤微生物群落结构与箭竹根际土壤微生物量碳、可溶性有机氮及箭竹生物量呈显著正相关。综上,干旱降低了箭竹根际土壤养分含量和微生物生物量,改变了箭竹根际土壤微生物群落结构,抑制了箭竹的生长;施磷能增加受旱箭竹根际土壤养分含量和微生物生物量,改善受旱箭竹根际土壤微生物群落结构,进而改善受旱箭竹的生长。     

The effect of vapor pressure deficit on maize transpiration response to a drying soil

A decline in plant transpiration has been widely observed to occur within a fairly stable range of threshold values of fraction transpirable soil water (FTSW), usually 0.3–0.4. However, the stability of this function has not been compared at various levels of atmospheric vapor pressure deficit (VPD). Soil hydraulic conductivity is likely to be involved in determining the threshold where water supply is limiting. Thus, it was hypothesized that at a high VPD resulting in increased transpiration rates, the FTSW threshold for the decline of transpiration rates as a result of drying soil would be increased. This study was undertaken in controlled environment chambers with two maize (Zea mays L.) hybrids (Pioneer Brand Hybrids `3165' and `3737') so as to subject plants to four VPD levels (1.1, 2.0, 2.9 and 3.6 kPa) during a soil drying experiment. In contrast to the original hypothesis, there was little ( 0.05 FTSW) change in the threshold FTSW in response to increased VPD for either hybrid. In fact, over the narrow 0.31–0.38 FTSW range observed, the two hybrids showed opposite trends in FTSW threshold as VPD increased. These results supported the view that the FTSW threshold for the decline in transpiration with drying soil is stable, showing little sensitivity to changes in VPD.     

夏玉米对土壤水分持续减少的响应及其转折点阈值分析

玉米是世界三大粮食作物之一,玉米生产在中国粮食安全与畜牧业发展中具有举足轻重的作用。干旱是夏玉米生产最主要气象灾害,及时准确地获取干旱信息对夏玉米安全生产至关重要。以夏玉米郑单958品种为材料,设置充分供水和拔节期开始土壤水分持续减少两种水分处理,研究夏玉米对土壤水分持续减少的响应及其转折点阈值,为夏玉米干旱识别与监测提供依据。结果表明,土壤水分持续减少10d后生理指标开始陆续受到胁迫,20d后生物量积累受到抑制,30d左右形态特征开始受到胁迫。夏玉米生理指标中最先受到胁迫的是顶端第1片完全展开叶的含水量和水势,生物量积累指标中为茎生物量,形态指标中为叶数。夏玉米顶端第1片完全展开叶的含水量或水势、茎生物量和叶数开始受到土壤干旱过程胁迫的时间阈值分别为11、21、27d,水分亏缺阈值分别为34、66、86mm,土壤相对湿度阈值分别为64%、56%和52%。表明夏玉米对土壤干旱过程的响应首先表现为生理特征变化、其次为生物量积累变化、最后为形态特征变化。研究结果可为客观辨识夏玉米干旱的发生发展及监测预警提供参考。     

Xylem ABA controls the stomatal conductance of field-grown maize subjected to soil compaction or soil drying

Stomatal conductance of individual leaves was measured in a maize field, together with leaf water potential, leaf turgor, xylem ABA concentration and leaf ABA concentration in the same leaves. Stomatal conductance showed a tight relationship with xylem ABA, but not with the current leaf water status or with the concentration of ABA in the bulk leaf. The relationship between stomatal conductance and xylem [ABA] was common for variations in xylem [ABA] linked to the decline with time of the soil water reserve, to simultaneous differences between plants grown on compacted, non-compacted and irrigated soil, and to plant-to-plant variability. Therefore, this relationship is unlikely to be fortuitous or due to synchronous variations. These results suggest that increased concentration of ABA in the xylem sap in response to stress can control the gas exchange of plants under field conditions.     

Non-hydraulic signals from maize roots in drying soil: inhibition of leaf elongation but not stomatal conductance

Conditions of soil drying and plant growth that lead to non-hydraulic inhibition of leaf elongation and stomatal conductance in maize (Zea mays L.) were investigated using plants grown with their root systems divided between two containers. The soil in one container was allowed to dry while the other container was kept well-watered. Soil drying resulted in a maximum 35% inhibition of leaf elongation rate which occurred during the light hours, with no measurable decline in leaf water potential (_w). Leaf area was 15% less than in control plants after 18 d of soil drying. The inhibition of elongation was observed only when the soil _w declined to below that of the leaves and, thus, the drying soil no longer contributed to transpiration. However, midday root _w in the dry container (-0.29 MPa) remained much higher than that of the surrounding soil (-1.0 MPa) after 15 d of drying, indicating that the roots in drying soil were rehydrated in the dark.To prove that the inhibition of leaf elongation was not caused by undetectable changes in leaf water status as a result of loss of half the watergathering capacity, one-half of the root system of control plants was excised. This treatment had no effect on leaf elongation or stomatal conductance. The inhibition of leaf elongation was also not explained by reductions in nutrient supply.Soil drying had no effect on stomatal conductance despite variations in the rate or extent of soild drying, light, humidity or nutrition. The results indicate that non-hydraulic inhibition of leaf elongation may act to conserve water as the soil dries before the occurrence of shoot water deficits.Symbol _w water potential Contribution from the Missouri Agricultural Experiment Station, Journal Series No. 10881     

Changes in plant–soil hydraulic pressure gradients of soybean in response to soil drying

Plant gas‐exchange response to drying soil in many instances tracks a common pattern when expressed as a function of fraction of transpirable soil water (FTSW). There is little decrease in gas exchange until FTSW decreases to a value in the range of about 0.3–0.45, then with further drying gas exchange declines approximately linearly. This unique pattern is hypothesised to reflect mainly changes in the water potential gradient between bulk soil and plant. The primary objective was to directly document the basis of this response by measuring the hydrostatic pressure gradient required in the soil to maintain leaf xylem at zero potential with decreasing FTSW. Pots in which soybean (Glycine max) plants were grown were placed in a pressure chamber and the pressure adjusted to maintain zero water potential in a leaf petiolule. These results showed a small, relatively constant hydrostatic pressure had to be applied to the soil to maintain zero leaf xylem water potential until FTSW decreased to approximately 0.3–0.45. Thereafter, the required hydrostatic pressure gradient increased as FTSW continued to decrease. Hydraulic conductance was calculated to be relatively stable early in the drying cycle, and then decrease as the soil dried to comparatively high FTSW of 0.5–0.7.     

Growth and gas exchange response to water shortage of a maize crop on different soil types

The effect of water shortage on growth and gas exchange of maize grown on sandy soil (SS) and clay soil was studied. The lower soil water content in the SS during vegetative growth stages did not affect plant height, above-ground biomass, and leaf area index (LAI). LAI reduction was observed on the SS during the reproductive stage due to early leaf senescence. Canopy and leaf gas exchanges, measured by eddy correlation technique and by a portable photosynthetic system, respectively, were affected by water stress and a greater reduction in net photosynthetic rate (A _N) and stomatal conductance (g _s) was observed on SS. Chlorophyll and carotenoids content was not affected by water shortage in either condition. Results support two main conclusions: (1) leaf photosynthetic capacity was unaffected by water stress, and (2) maize effectively endured water shortage during the vegetative growth stage.     

Soybean leaf growth and gas exchange response to drought under carbon dioxide enrichment

This study was conducted to determine the response in leaf growth and gas exchange of soybean (Glycine max Merr.) to the combined effects of water deficits and carbon dioxide (CO₂) enrichment. Plants grown in pots were allowed to develop initially in a glasshouse under ambient CO₂ and well-watered conditions. Four-week old plants were transferred into two different glasshouses with either ambient (360 μmol mol^-1) or elevated (700 μmol mol^-1) CO₂. Following a 2-day acclimation period, the soil of the drought-stressed pots was allowed to dry slowly over a 2-week period. The stressed pots were watered daily so that the soil dried at an equivalent rate under the two CO₂ levels. Elevated [CO₂] decreased water loss rate and increased leaf area development and photosynthetic rate under both well-watered and drought-stressed conditions. There was, however, no significant effect of [CO₂] in the response relative to soil water content of normalized leaf gas exchange and leaf area. The drought response based on soil water content for transpiration, leaf area, and photosynthesis provide an effective method for describing the responses of soybean physiological processes to the available soil water, independent of [CO₂].     

Precipitation regulates plant gas exchange and its long-term response to climate change in a temperate grassland

Aims Climate change largely impacts ecosystem carbon and water cycles by changing plant gas exchange, which may further cause positive or negative feedback to global climate change. However, long-term global change manipulative experiments are seldom conducted to reveal plant ecophysiological responses to climatic warming and altered precipitation regimes.Methods An 8-year field experiment with both warming and increased precipitation was conducted in a temperate grassland in northern China. We measured leaf gas exchange rates (including plant photosynthesis, transpiration and instantaneous water use efficiency [WUE]) of two dominant plant species (Stipa sareptana var. krylovii and Agropyron cristatum) from 2005 to 2012 (except 2006 and 2010) and those of other six species from 2011 to 2012.Important findings Increased precipitation significantly stimulated plant photosynthetic rates (A) by 29.5% and 19.9% and transpiration rates (E) by 42.2% and 51.2% for both dominant species S. sareptana var. krylovii and A. cristatum, respectively, across the 8 years. Similarly, A and E of the six plant functional types were all stimulated by increased precipitation in 2011 and 2012. As the balance of A and E, the instantaneous WUEs of different plant species had species-specific responses to increased precipitation. In contrast, neither warming nor its interaction with increased precipitation significantly affected plant leaf gas exchange rates. Furthermore, A and E of the two dominant species and their response magnitudes to water treatments positively correlated with rainfall amount in July across years. We did not find any significant difference between the short-term versus long-term responses of plant photosynthesis, suggesting the flexibility of leaf gas exchange under climate change. The results suggest that changing precipitation rather than global warming plays a prominent role in determining production of this grassland in the context of climate change.     

Wetting and drying cycles in the maize rhizosphere under controlled conditions. Mechanics of the root-adhering soil

Mechanical properties of the topsoil (sandy Podsol and silty Luvisol, FAO) adhering to maize (Zea mays L.) roots and its bulk soil counterpart were studied as a function of soil texture and final soil water suction at harvest, with three soil water suction values of approximately 30, 50 and 60 kPa. Two scales of observation were also selected: the whole soil:root system and the root-adhering soil aggregates. Three methods were used to characterize the stability of the soil:root system: mechanical shaking in air, and dispersion by low-power ultrasonication, with or without preliminary immersion of the soil:root system in water. Soil disruption kinetics, which were fitted with first-order kinetics equations, were analyzed and discussed. For example, silty soil ultrasonication kinetics, without preliminary water-immersion, could be divided into two parts: the first faster part, which was characterized by a mean rate K value of 6.8–7.2 mJ^-1, is attributed to soil slaking, whereas the second slower part, which was characterized by a mean rate K value of 1.5–1.6 mJ^-1, was attributed to the rupture of the `firmly root-adhering soil' from the roots. A clear plant effect was observed for both aggregate tensile strength and friability, with higher aggregate strength for the root-adhering silty soil (450–500 kPa) than for its bulk silty soil counterpart (410–420 kPa), and lower friability (coefficient of variation of the aggregate strength) for the root-adhering silty soil (e.g. 67% at a soil water suction value of 30 kPa) than for its bulk silty soil counterpart (e.g. 49% at asoil water suction value of 30 kPa). These effects were attributed to root exudation, which was significantly higher for the driest silty topsoil than for the wetter ones. In conclusion, the mechanical properties of the silty topsoil adhering to the maize roots are attributed to both physical and biological interactions occurring in the maize rhizosphere. This revised version was published online in June 2006 with corrections to the Cover Date.