控释尿素水氮耦合对夏玉米产量和光合特性的影响

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李广浩, 赵斌, 董树亭, 刘鹏, 张吉旺, 何在菊. 控释尿素水氮耦合对夏玉米产量和光合特性的影响. 作物学报, 2015, 41(9): 1406-1415
LI Guang-Hao, ZHAO Bin, DONG Shu-Ting, LIU Peng, ZHANG Ji-Wang, HE Zai-Ju. Effects of Coupling Controlled Release Urea with Water on Yield and Photosynthetic Characteristics in Summer Maize. ACTA AGRONOMICA SINICA, 2015, 41(9): 1406-1415 复制到剪切板

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作物学报编辑部

控释尿素水氮耦合对夏玉米产量和光合特性的影响

李广浩, 赵斌^*, 董树亭^*, 刘鹏, 张吉旺, 何在菊

山东农业大学农学院 / 作物生物学国家重点实验室, 山东泰安271018

^* 通讯作者(Corresponding authors): 赵斌, E-mail:zhaobin@sdau.edu.cn, Tel: 0538-8245838; 董树亭, E-mail:stdong@sdau.edu.cn

第一作者联系方式: 李广浩, E-mail:guanghaoli@126.com, Tel: 15275381215

收稿日期:2014-12-25 接受日期:2015-06-01 网络出版日期:2015-06-23

基金:本研究由国家自然科学基金项目(31171497), 国家公益性行业(农业)科研专项(201203096), 国家现代农业产业技术体系建设专项(CARS-02-20), 国家重点基础研究发展计划(973计划)项目(2011CB100105), 国家粮食丰产科技工程项目(2011BAD16B09), 山东省良种工程重大课题(鲁科农字[2011]186号), 国家自然科学基金青年基金项目(31301274)和山东省高等学校科技计划项目(J13LF08)资助

摘要

采用旱棚盆栽试验, 以郑单958为材料设置3个水分水平(正常水分W3、轻度水分胁迫W2、重度水分胁迫W1)和高氮N3 (施纯氮315 kg hm^-2)、中氮N2 (施纯氮210 kg hm^-2)、低氮N1 (施纯氮105 kg hm^-2)、不施氮N0四个控释尿素施氮水平, 探讨控释尿素水氮耦合对夏玉米产量和光合特性的影响。结果表明, 控释尿素水氮耦合对夏玉米产量和光合特性具有显著影响。相同水分条件下, 夏玉米产量随施氮量增加而增加, W1条件下增产13.17%~20.96%, W2条件下增产13.93%~32.48%, W3条件下增产14.37%~21.83%。相同施氮水平下, 产量也随水分增加而增加, W2N3、W3N2和W3N3的产量在所有处理中较高。水氮耦合对夏玉米穗位叶净光合速率的影响显著, W1条件下N3、N2和N1处理间差异不显著, 均显著高于N0, W2、W3各施氮处理的净光合速率随施氮量增加而增加, W3各处理的平均净光合速率高于其他2个水分处理, W2N3比W3N3和W3N2前期略低, 后期无显著差异。水氮耦合效应能有效减缓穗位叶的实际光化学效率 Φ_PSII、叶片光化学猝灭系数 q_P以及PSII反应中心的最大光能转换效率的下降速率, 提高光能利用率。控释尿素水氮耦合能有效提高夏玉米花后穗位叶净光合速率, 保证籽粒对营养物质的需求, 提高穗位叶实际和最大光化学效率, 从而提高夏玉米的产量, 产量构成因素中增加千粒重和穗粒数的优势较大。综合产量与光合特性、荧光特性的表现, 在田间持水量为75%±5%的土壤条件下, 控释尿素施氮量以纯氮210 kg hm^-2为最佳; 在田间持水量为55%±5%的土壤条件下, 控释尿素施氮量以纯氮315 kg hm^-2为宜。

关键词: 夏玉米; 控释尿素; 水氮耦合; 产量; 光合特性

Effects of Coupling Controlled Release Urea with Water on Yield and Photosynthetic Characteristics in Summer Maize

LI Guang-Hao, ZHAO Bin^*, DONG Shu-Ting^*, LIU Peng, ZHANG Ji-Wang, HE Zai-Ju

Agronomy College of Shandong Agricultural University / State Key Laboratory of Crop Biology, Tai’an 271018, China

Abstract

A experiment using Zhengdan 958 (a summer maize cultivar planted widely in China), with treatments of three water levels (normal water W3, mild water stress W2, severe water stress W1) and four amounts of controlled release urea nitrogen (N) (N3 was 315 kg ha^-1, N2 was 210 kg ha^-1, N1 was 105 kg ha^-1 and N0 was no nitrogen) was carried out under the waterproof shed and potting conditions. The results showed there was significant influence of coupling water with controlled release urea on yield and photosynthetic characteristics in summer maize. Under the condition of same soil moisture, the yield showed a increasing trend with the amounts of N application increased. The yield increased by 13.17%-20.96% under severe water stress (W1), by 13.93%-32.48% under mild water stress (W2), and by 14.37%-21.83% under the condition of normal water (W3). Under the same amounts of N application, the yield was also improved with the increase of the soil moisture. The yields of W2N3, W3N2, and W3N3 were the highest among these treatments. The coupling effects of water and controlled release urea were significant on the photosynthetic rate in ear leaf. The photosynthetic rates of N3, N2, and N1 had no significant differences in W1, and were higher than these of N0 significantly. The photosynthetic rate of W2 and W3 treatments increased with the increase of N fertilizer application, the average photosynthetic rate in W3 was higher than that of W2 and W1, that of W2N3 was slightly lower than that of W3N3 and W3N2 at early flowering, and no significant difference in the late stage. The coupling effects of water and controlled release urea nitrogen could effectively improve the efficiency of light energy utilization and slow down the decline rates of the actual photochemical efficiency of PSII, coefficient of photochemical quenching ( q_P) of leaf and PSII reaction center of light conversion. High photosynthetic rate, actual and maximal photochemical efficiency of ear leaf after flowering in summer maize ensured the nutrients needs of grain filling, thus increased the yield. Higher yield was obtained due to the significant increase of kernels per ear and thousand seed weight among yield components. Under this experimental condition, controlled release urea N application rate of 210 kg ha^-1 was the best treatment as the soil moisture content was 75%±5% of field capacity. We suggested that coupling controlled release urea N application rate of 315 kg ha^-1 with soil moisture content of 55%±5% of field capacity is optimum.

Keyword: Summer maize; Controlled release urea; Water and nitrogen coupling; Yield; Photosynthetic characteristics

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玉米是我国第一大作物, 在粮食生产中举足轻重。随着人口的增长及耕地面积的减少, 玉米在作物生产中的地位越来越重要。当前, 品种改良和化肥投入是玉米总产增长的贡献因子中最主要的因素^{[1, 2, 3, 4]}。据联合国粮农组织估计, 在发展中国家的粮食增产中, 有55%来自化肥的作用^[5]。受社会经济条件的制约, 目前中国玉米生产中一次性施肥(肥料全部做基肥, 又称“ 一炮轰” )的农户占很大比例, 米国华课题组2009年在吉林梨树县调查发现, 当地一次性施肥比例高达2/3以上^[6], 黄淮海玉米主产区亦如此。如何协调肥料投入与产量之间的矛盾已引起广泛重视, 氮素释放速率与作物需肥规律基本一致、一次性基施满足生产需要、同时能有效提高生产效率、减少人工投入的缓控释肥的研发与应用被寄予厚望^{[7, 8, 9, 10, 11, 12]}。明确不同水分条件下控释肥的适宜施用量有助于黄淮海地区夏玉米专用控释肥的合理应用。水氮供应是调控作物生长和发育的重要手段, 通过合理的水氮运筹调节作物光合特性及光合产物的积累和分配, 是实现作物高产的重要措施之一^[13]。适时适度减少水分供应不会降低作物光合速率, 却显著减少作物蒸腾失水, 从而提高作物水分利用效率^[14]。大量研究和实践表明, 作物的光合作用及产量依赖于土壤中的水肥变化。徐世昌等研究表明水分胁迫导致玉米光合性能及产量的降低^[15]。在土壤干旱状况下增施氮肥, 或者在低肥条件下增加灌水量, 均可显著提高玉米产量; 在节水节肥条件下, 通过氮水耦合也可以实现玉米高产^[16]。李潮海等^[17]指出, 施肥能显著增加玉米的光合速率, 并且随着施肥量的增加后期高光合速率持续期延长。夏玉米的整个生育期内, 气温较高, 植株高大, 给追肥带来很大困难, 需要消耗大量的劳动力, 一些地区出现减少追肥次数的现象, 导致玉米不同程度减产, 给农民造成损失。控释尿素成为最好的解决方法, 然而不同生态区的降水差异很大, 不同的降水量必然影响控释尿素的释放, 目前国内在玉米上有关控释肥与水分互作的应用研究报道尚少, 因此探索不同水分条件下控释尿素的适宜施用量非常必要。本试验重点研究控释尿素与水分耦合对夏玉米产量和光合特性的影响, 从光合特性、荧光特性及产量方面, 深入探讨水氮耦合对夏玉米生长发育的调控机制, 为黄淮海地区夏玉米专用控释肥的应用提供科学依据, 为节水农业提供理论基础。

1 材料与方法

1.1 试验地状况

山东农业大学黄淮海区域玉米技术创新中心(36° 10′ N, 117° 09′ E)地处黄淮海平原, 属于半湿润暖温带大陆性季风气候区。试验用土壤为中壤土, 耕层0~20 cm土壤pH 6.1, 含有机质19.7 g kg^-1、全氮1.1 g kg^-1、碱解氮124.4 mg kg^-1、速效磷45.2 mg kg^-1、速效钾81.8 mg kg^-1, 土壤田间持水量为21.1%, 土壤容重1.5 g cm^-3。

1.2 试验设计

2012年和2013年采用防雨旱棚(只在雨天使用), 盆栽试验泥塑盆直径35 cm, 高45 cm, 试验玉米品种为郑单958, 设置3个水分处理, W1的水分含量为田间持水量的35%± 5%; W2的水分含量为田间持水量的55%± 5%; W3的水分含量为田间持水量的75%± 5%。用TDR土壤水分仪控制土壤水分, 每天上午和傍晚各测定一次以确定每天的浇水量。采用山东农业大学资源与环境学院研制的包膜控释尿素(氮素含量42%, 控释周期为3个月), 设置4个施氮处理, N0为施纯氮0 kg hm^-2, 每盆0 g; N1为施纯氮105 kg hm^-2, 每盆1.56 g; N2为施纯氮210 kg hm^-2, 每盆3.12 g; N3为施纯氮315 kg hm^-2, 每盆4.68 g。

每盆装风干土(过4 mm筛孔) 35 kg, 每个处理均施P₂O₅ 105 kg hm^-2, K₂O为210 kg hm^-2, 即每盆施P₂O₅ 1.56 g, K₂O 3.12 g。共12个处理, 每盆一株, 每个处理15盆, 总共180盆。6月18日播种, 播种前灌水至田间持水量, 以保证全苗、壮苗。所有氮肥、磷肥和钾肥均于播种前一次性作底肥施用。

1.3 试验取样

从开花期开始每隔10 d取1次样, 选每处理生长一致的植株1株, 取穗位叶放入液氮罐带回室内, 置-40℃冰箱保存以测定叶绿素含量。成熟期收获(苞叶枯黄、乳线消失、黑层出现)各处理, 分别取5穗晒干, 室内考种, 调查穗长、穗粗、秃顶长、穗行数、行粒数、千粒重。单株产量(g)=穗粒数× 千粒重(g)/1000× [1-含水量(%)]/(1-14%)。

1.4 测定指标

各生育时期取样时, 测量叶面积(定株测量)。

采用Arnon法95%乙醇提取, UV-2800型紫外可见分光光度计测定和计算光合色素含量。

采用美国LI-COR公司生产的LI-6400光合测定系统, 以开放式光路, 于晴天10:00— 14:00测定光合速率, 测定部位为穗位叶中部上表面, 于开花期开始, 每隔15 d测定一次。

采用德国Walz公司生产的PAM脉冲调制式荧光仪测定叶绿素荧光动力学参数。参考张守仁^[18]的方法计算叶绿素荧光动力学参数。

1.5 统计分析

采用Microsoft Excel 2003处理数据。用DPS 7.05软件统计分析, 在0.05水平进行显著性检验(LSD, least significant difference test)。

2 结果与分析

2.1 控释尿素水氮耦合对夏玉米产量及构成因素的影响

从表1可以看出, 控释尿素与水分对夏玉米产量影响显著。相同水分条件下, 夏玉米的产量随着施氮量的增加呈增加趋势, 其中重度水分胁迫(W1)下比N0增产13.17%~20.96%; 轻度水分胁迫(W2)下增产幅度相对较大, 较N0增产13.93%~32.48%; 而正常水分(W3)条件下比N0增产14.37%~ 21.83%。相同氮肥水平, 不同水分条件, 各处理产量也呈现随着水分的增加而增加的趋势, N0情况下, W3和W2分别比W1增产36.23%和40.07%; N1水平下, W3和W2分别增产37.07%和56.13%; N2水平下, 分别增产39.28%和63.75%; N3水平下, 分别增产49.12%和54.83%。从表1还可看出, W3水分条件下, N2与N3产量无显著差异, 说明N2处理肥量适宜, N3处理过量; 在W2水平下, 各肥料处理中N3产量最高, 差异显著。经显著性检验(表2), 水分为主效应, 控释尿素水氮耦合效应达极显著水平(P< 0.01)。在产量构成因素中, 千粒重和穗粒数是影响夏玉米产量的最关键因素, 产量高的处理其千粒重与穗粒数都显著高于产量低的处理, 并达显著水平。

表1 控释尿素水氮耦合对夏玉米产量及其构成因素的影响 Table 1 Effects of coupling controlled release urea with water on yield and yield components of summer maize

表2 控释尿素水氮耦合对夏玉米产量影响显著性检验 Table 2 Significant analysis of effects of coupling controlled release urea with water on yield of summer maize

2.2 控释尿素水氮耦合对夏玉米光合特性的影响

2.2.1 穗位叶净光合速率(P_n) 从图1可以看出, 控释尿素水氮耦合对玉米穗位叶净光合速率的影响显著, 花后随着时间的推进, 各处理的光合速率持续下降, 前期下降相对缓慢, 后期下降较快, 到成熟期降至最低。开花期正常水分W3处理的光合速率最高, W2次之, W1最低; W2各施肥处理随着施氮量的增加光合速率也随着增加, W1各施氮处理之间差异不显著, 均显著高于N0, W3条件下, 前期表现为N2和N3最高, N1次之, 到后期各施肥处理差异不显著, 均显著高于N0, 说明施氮显著提高玉米花后净光合速率, 有利于产量的提高。从开花到成熟, W3的各处理的平均光合值显著高于其他2个水分处理, W2N3比W3N3和W3N2前期略低, 后期三者无显著差异。经显著性检验(表3), 夏玉米光合速率的水氮耦合效应达显著水平, 后期水分的效应高于氮肥的效应。

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	图1 夏玉米穗位叶净光合速率变化缩写同表1。Fig. 1 Changes of photosynthetic rate in summer maize ear leafAbbreviations are the same as those given in Table 1.

表3 控释尿素水氮耦合对夏玉米穗位叶净光合速率P_n影响的显著性检验(F值) Table 3 Significant analysis of effects of coupling controlled release urea with water on photosynthetic rate in summer maize ear leaf

2.2.2 气孔导度(G_s) 气孔导度控制着植物体与外界环境的水、气交换, 对植物光合作用有很大的影响。从表4可以看出, 夏玉米穗位叶气孔导度(G_s)随着生育进程而逐渐下降。开花期, 相同施氮量情况下, 正常水分W3处理的G_s显著高于W2和W1处理, W2显著高于W1。同一水分条件下, 呈现随着施氮量的增加而升高的趋势, W1条件下, 各处理分别比N0平均增加14.85%、11.41%和19.76%; W2条件下各处理分别比N0增加6.48%、17.91%和25.65%; W3条件下分别比N0增加11.62%、27.43%和25.93%。W3N2与W3N3、W2N2与W2N3、W1N2与W1N3无差异, 从开花到成熟都表现出相同的规律。成熟期各处理的G_s表现为W3> W2> W1。

表4 控释尿素水氮耦合对夏玉米穗位叶气孔导度G_s的影响 Table 4 Effects of coupling controlled release urea with water on G_s in summer maize ear leaf (mmol m^-2 s^-1)

2.2.3 夏玉米开花后穗位叶叶绿素总含量的变化

由图2可以看出, 随生育进程叶绿素总含量不断下降。开花期叶绿素总含量最高, 到花后10 d下降平缓, W1水平下有所升高, 之后下降迅速, 到成熟期达到最低。在W1水平下, 前期叶绿素含量表现为N3> N2> N1> N0, 后期各处理差异不显著; W2水平下, 一直表现为N3> N2> N1> N0, 差异显著, 成熟期N2与N1无显著差异; 在W3水平下, N2略高于N3, 差异不显著, 到成熟期二者无差异, 都显著高于N1和N0。相同施肥量N3情况下, W3略高于W2, 差异不显著, 二者都显著高于W1; 在N2、N1和N0条件下, 均表现出W3> W2> W1, 差异达显著水平。

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	图2 夏玉米开花后穗位叶叶绿素含量的变化缩写同表1。Fig. 2 Changes of chlorophyll content in summer maize ear leaf after anthesisAbbreviations are the same as those given in Table 1.

2.2.4 叶面积指数(LAI)的变化随着生育期的推迟, 各处理花后叶面积指数均呈现逐渐降低的变化(图3)。在W3水平下, 从开花到花后20 d, 各处理叶面积指数下降相对平缓, 之后下降迅速, 但叶面积指数保持较高; 在W2水平下, 各施氮处理前期下降也相对平缓, 后期迅速下降, 其总体叶面积指数要低于W3; W1水平下, 叶面积指数从花后10 d就开始下降, 花后30 d到40 d急剧下降, 叶面积指数很低。相同水分条件下, W1的各施氮处理差异不显著, W2的各施氮处理中N3显著高于N2和N1, N2和N1差异不显著, W3的各施氮处理中N3和N2差异不显著, 两处理显著高于N1, 所有施氮处理均显著高于相同水分的N0。

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	图3 控释尿素水氮耦合对夏玉米花后叶面积指数的影响缩写同表1。Fig. 3 Effects of coupling controlled release urea with water on leaf area index after anthesis of summer maizeAbbreviations are the same as those given in Table 1.

2.3 控释尿素水氮耦合对夏玉米穗位叶荧光特性的影响

2.3.1 穗位叶实际光化学效率Φ _PSII变化从图4可以看出, 夏玉米穗位叶的实际光化学效率Φ _PSII随着生育进程而逐渐下降。W1水平下, 各施氮处理下降迅速, W2次之, 而W3各处理下降相对缓慢。W1水平下, 一直表现为N3> N2> N1> N0, N3显著高于N2与N1, N2与N1差异不显著; W2水平下, 一直表现为N3> N2> N1> N0, N3显著高于N2与N1, N2与N1前期差异显著, 后期无显著差异; W3水平下, N3与N2无显著差异, 但都显著高于N1和N0。相同氮肥条件下, Φ _PSII随着水分的降低而下降, 差异显著。但相同水分条件下N2与N3差异不显著。

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	图4 夏玉米穗位叶实际光化学效率变化缩写同表1。Fig. 4 Changes of Φ _PSII in summer maize ear leafAbbreviations are the same as those given in Table 1.

2.3.2 穗位叶最大光化学效率F_v/F_m变化花后夏玉米穗位叶的最大光化学效率随着生育进程而逐渐下降, 表现为W1> W2> W3。穗位叶最大光化学效率在3个水分条件下都表现为随施氮量增加而升高的趋势。W1水平下, N3显著高于N2与N1, N2与N1间差异不显著; W2水平下, N3与N2前期差异显著, 后期差异不显著, 都显著高于N1和N0; W3水平下, N3与N2一直无显著差异, 但都显著高于N1和N0。相同氮肥条件下, F_v/F_m随着水分的降低而下降, 差异显著。但W3N2、W3N3与W2N3差异不显著, W3N1与W2N2差异不显著。

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	图5 夏玉米穗位叶最大光化学效率变化缩写同表1。Fig. 5 Changes of F_v/F_m in summer maize ear leafAbbreviations are the same as those given in Table 1.

2.3.3 光化学猝灭与非光化学猝灭系数从表5可以看出, 叶片的光化学猝灭系数(q_P)随生育进程而下降, 非光化学猝灭系数(NPQ)则相反, 呈不断上升的趋势。相同水分条件下, 随着施氮量的增加, q_P明显增加, 而NPQ显著下降。W1条件下, 各处理的q_P比N0平均增加6.75%、11.01%和11.75%, 各处理的NPQ比N0平均降低23.89%、26.97%和28.25%; W2条件下, 各处理的q_P比N0平均增加6.60%、11.53%和13.97%, 各处理的NPQ比N0平均降低30.39%、34.69%和36.95%; W3条件下, 各处理的q_P比N0平均增加10.03%、13.63%和12.92%, 各处理的NPQ比N0平均降低30.15%、33.82%和33.58%。相同施氮量条件下, 各水分处理的q_P表现为W3> W2> W1, 差异显著。对NPQ来说, 各水分处理差异不显著, 尤其是W2与W3, 无明显差异。

表5 控释尿素水氮耦合对夏玉米穗位叶光化学猝灭和非光化学猝灭系数的影响 Table 5 Effects of coupling controlled release urea with water on q_P and NPQ in summer maize ear leaf

3 讨论

3.1 控释尿素水氮耦合对夏玉米光合特性的影响

作物产量的90%以上来自光合作用^[19], 光合作用是玉米干物质生产的基础, 其功能直接影响玉米籽粒产量。水氮对作物光合作用的影响是多方面的。宋凤斌等研究认为, 水分胁迫下玉米叶片光合速率和气孔导度随胁迫增强而下降, 而在重度胁迫下光合速率显著下降, 水分胁迫使玉米叶片的叶绿素含量降低^[20]。本试验结果表明, 随水分含量的增加光合速率、气孔导度和穗位叶叶绿素含量都表现出升高的趋势, 与宋凤斌等在玉米光合特性上的研究结果一致, 增施氮肥可以减缓水分胁迫对夏玉米光合特性的不利影响。常敬礼等研究表明氮肥提高了穗位叶叶片的净光合速率和叶绿素含量, 并且施氮肥越多, 各项光合参数值的增加幅度越大^{[21, 22, 23]}, 李潮海等^[17]指出, 施肥可以改善叶肉细胞的光合能力, 提高生育后期叶片的光合强度, 延长高光合持续期。本试验相同水分条件下随控释尿素增加净光合速率和叶绿素含量也有相同趋势, 但是W3N2与W3N3无显著差异, 表明在W3条件下N3处理比N2处理氮肥量增加, 而光合特性没有增强。本试验结果表明, 水氮对夏玉米光合特性影响的耦合效应达到了极显著的水平, 水分为主效应, 控释尿素为次效应, W2N3、W3N2和W3N3处理的花后穗位叶叶绿素含量相对于其他处理一直保持较高水平, 这为其保持较高的净光合速率、气孔导度和细胞间隙CO₂浓度奠定了基础, 提高了夏玉米的光合能力, 进而提高了产量。生育期内叶面积指数的变化强度和幅度是植株叶片生长发育和衰老的直接反映, W2和W3水分条件下, 施氮有效缓解了叶面积指数的下降速率, 延缓了叶片衰老, 且W2N3、W3N2和W3N3处理之间差异不显著, 控释尿素与水分耦合效应在花后20 d内可以有效缓解叶面积指数下降的速率, 提高光合速率, 为夏玉米高产提供保障。

3.2 控释尿素水氮耦合对夏玉米荧光特性的影响

通过叶绿素荧光可快速检测完整植株在水分胁迫下光合作用的真实行为, 用来评价光合机构的功能和环境胁迫的影响^[24]。荧光参数q_P反映PSII反应中心天线色素吸收的光能用于光化学电子传递率和PSII反应中心的开放程度, q_P愈大, PSII的电子传递活性愈大; NPQ反映PSII天线色素吸收的光能不能用于光合电子传递而以热的形式耗散的部分, 这种热耗散可以避免过量光能对光合结构的破坏。齐华等^[24]研究表明水分胁迫使玉米叶片光合性能减弱, PSII反应中心开放部分的比例减少, 光合电子传递能力下降, PSII潜在活性受到抑制, F_v/F_m、F_m和F_v降低, F_o升高, q_P降低, NPQ升高, 王建程等^[25]研究发现提高施肥水平有利于减轻水分胁迫对光合中心的伤害, 高肥有助于提高玉米PSII光化学效率。本试验结果表明, 相同水分条件下, 施氮量的增加能够显著提高叶片的Φ _PSII和PSII反应中心的最大光能转换效率; 相同氮肥条件下, Φ _PSII和叶片的最大光化学效率表现为W3> W2> W1, 对水分效应敏感, 这与李耕等^[26]运用JIP-test分析研究干旱胁迫对玉米叶片光系统活性影响的结果一致。随着施氮量的增加, q_P明显增加, 而NPQ显著下降, 表明适当施用氮肥可以提高叶片吸收光能的光化学利用, 而减弱非光化学利用。相同施氮量条件下, q_P表现为W3> W2> W1, 对水分效应敏感。对NPQ来说, 各水分处理差异不显著, 尤其是W2与W3, 表明对水分处理不敏感; 但是W2条件下的N1与N2差异不显著, 表明适当增施氮肥可以缓解轻度胁迫对夏玉米NPQ的影响, W3条件下的N2与N3差异不显著, 过量增加氮肥不能提高叶片的Φ _PSII和PSII反应中心的最大光能转换效率, 这是控释尿素与水分耦合效应的结果。

3.3 控释尿素水氮耦合对夏玉米产量的影响

前人研究表明, 相同水分条件下, 作物的产量随着施氮量的增加呈增加趋势; 相同氮肥水平条件下, 产量随水分增加而增加^{[27, 28, 29, 30, 31]}, 这与本试验结果一致。邵国庆等^[32]研究表明, 氮素和水分在玉米产量上存在显著的正耦合效应。本研究表明, 控释尿素水氮耦合对夏玉米产量具显著影响且存在互作效应, 水分为主效应, 控释尿素为次效应。重度水分胁迫下, 夏玉米穗位叶的叶绿素含量降低, 光合速率和气孔导度下降, Φ _PSII和PSII反应中心的最大光能转换效率下降, 产量降低, 增施氮肥对胁迫有所缓解, 产量随着施氮量的增加呈增加趋势; 轻度水分胁迫下N3的产量明显高于其他处理, 与正常水分条件下N2和N3处理差异不显著, 表明W2N3处理水氮耦合效应极显著, 有效缓解了轻度水分胁迫对穗位叶光合能力的影响, 增产最显著。正常水分条件下N2与N3处理产量无显著差异, N2处理增产最大, 表明在N2基础上继续增施氮肥, 其穗位叶光合能力不再继续提高, 增产效果不显著, 其原因是控释尿素与水分耦合效应的结果, 控释尿素在整个生育期都保持较高的氮素水平, 氮素释放缓慢而平稳, 能够保证玉米后期的氮素供应。在W2N3处理下, 施氮有效缓解了水分胁迫, 使产量提高。W3N3处理的产量没有继续增加, 表明其施氮量可能过量, 这与赵斌等^{[9, 11]}和韩晓日等^[33]在控释尿素上的研究结果一致。不同水分条件下控释尿素在土壤中的迁移影响夏玉米产量的机制有待进一步研究。

4 结论

控释尿素与水分在提高夏玉米的光合特性方面具有显著的耦合效应, 水分为主效应, 控释尿素为次效应, 合理的水氮运筹能够有效提高夏玉米穗位叶的叶绿素含量, 增强花后穗位叶的光合和荧光特性, 促进光合产物向籽粒的运转与分配, 进而提高产量, 特别是能提高产量构成因素中的千粒重和穗粒数。控释尿素施纯氮210 kg hm^-2和田间持水量为75%± 5%的条件下耦合效应最佳, 在田间持水量为55%± 5%条件下, 控释尿素施纯氮以315 kg hm^-2为宜。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。The authors have declared that no competing interests exist.

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[24]	齐华, 白向历, 孙世贤, 刘明, 杨国航, 丛雪. 水分胁迫对玉米叶绿素荧光特性的影响. 华北农学报, 2009, 24(3): 102-106 Qi H, Bai X L, Sun S X, Liu M, Yang G H, Cong X. Effects of water stress on chlorophyll fluorescence parameters of maize. Acta Agric Boreali-Sin, 2009, 24(3): 102-106 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.289]
[25]	王建程, 严昌荣, 卜玉山. 不同水分与养分水平对玉米叶绿素荧光特性的影响. 中国农业气象, 2005, 26(2): 95-98 Wang J C, Yan C R, Pu Y S. Effects of vary soil moisture and fertility levels on chlorophyll fluorescence characteristics in maize. Chin J Agrometeorol, 2005, 26(2): 95-98 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.835]
[26]	李耕, 高辉远, 赵斌, 董树亭, 张吉旺, 杨吉顺, 王敬锋, 刘鹏. 灌浆期干旱胁迫对玉米叶片光系统活性的影响. 作物学报, 2009, 35: 1916-1922 Li G, Gao H Y, Zhao B, Dong S T, Zhang J W, Yang J S, Wang J F, Liu P. Effects of drought stress on activity of photo systems in leaves of maize at grain filling stage. Acta Agron Sin, 2009, 35: 1916-1922 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.681]
[27]	张秋英, 刘晓冰, 金剑, 李艳华, 王光华. 水肥耦合对玉米光合特性及产量的影响. 玉米科学, 2001, 9(2): 64-67 Zhang Q Y, Liu X B, Jin J, Li Y H, Wang G H. Effect of water and fertilizer coupling on photosynthetic characteristic and yield in corn. J Maize Sci, 2001, 9(2): 64-67 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.125]
[28]	武荣, 李援农. 水氮耦合对冬小麦根系分布和根冠比及产量的影响. 南方农业学报, 2013, 44: 963-967 Wu R, Li Y N. Effect of water and nitrogen fertilizer coupling on root distribution, root/shoot ratio and yield of winter wheat. J Southern Agric, 2013, 44: 963-967 (in Chinese) [本文引用:1]
[29]	魏永霞, 何双红, 魏永华. 控制灌溉条件下水肥耦合对水稻产量及其构成因子的影响. 灌溉排水学报, 2010, 29(5): 98-102 Wei Y X, He S H, Wei Y H. Influence of water fertilizer coupling on rice yield and compensation under the condition of water control irrigation. J Irrig Drain, 2010, 29(5): 98-102 (in Chinese with English abstract) [本文引用:1] [CJCR: 0.548]
[30]	温利利, 刘文智, 李淑文, 文宏达. 水肥耦合对夏玉米生物学特性和产量的影响. 河北农业大学学报, 2012, 35(3): 18-23 Wen L L, Liu W Z, Li S W, Wen H D. Effects of water and fertilizer coupling on biological characteristics and yields of summer maize. J Agric Univ Hebei, 2012, 35(3): 18-23 (in Chinese with English abstract) [本文引用:1] [CJCR: 0.567]
[31]	张学军, 赵营, 陈晓群, 王晓彬, 蔡典雄. 不同水氮供应对水稻产量、吸氮量及水氮利用效率的影响. 中国农学通报, 2010, 26(4): 126-131 Zhang X J, Zhao Y, Chen X Q, Wang X B, Cai D X. Effect of different water and N supply on the yield, N uptake and water/N use efficiency of rice. Chin Agric Sci Bull, 2010, 26(4): 126-131 (in Chinese with English abstract) [本文引用:1]
[32]	邵国庆, 李增嘉, 宁堂原, 张民, 江晓东, 王芸, 赵建波, 吕美蓉, 赵杰. 不同水分条件下常规尿素和控释尿素对玉米根冠生长及产量的影响. 作物学报, 2009, 35: 118-123 Shao G Q, Li Z J, Ning T Y, Zhang M, Jiang X D, Wang Y, Zhao J B, Lü M R, Zhao J. Effects of normal urea and release- controlled urea on root and shoot growth and yield of maize in different water conditions. Acta Agron Sin, 2009, 35: 118-123 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.681]
[33]	韩晓日. 新型缓/控释肥料研究现状与展望. 沈阳农业大学学报, 2006, 37: 3-8 Han X R. Current situation and prospects of new type slow and controlled releaseing fertilizers. J Shenyang Agric Univ, 2006, 37: 3-8 (in Chinese with English abstract) [本文引用:1] [CJCR: 0.556]

1991

5.021

0.0

... 当前, 品种改良和化肥投入是玉米总产增长的贡献因子中最主要的因素^[1,2,3,4] ...

1992

0.368

0.0

... 当前, 品种改良和化肥投入是玉米总产增长的贡献因子中最主要的因素^[1,2,3,4] ...

2002

2.608

0.0

... 当前, 品种改良和化肥投入是玉米总产增长的贡献因子中最主要的因素^[1,2,3,4] ...

2006

0.368

0.0

... 当前, 品种改良和化肥投入是玉米总产增长的贡献因子中最主要的因素^[1,2,3,4] ...

2001

0.0

... 据联合国粮农组织估计, 在发展中国家的粮食增产中, 有55%来自化肥的作用^[5] ...

2010

1.512

0.692

... )的农户占很大比例, 米国华课题组2009年在吉林梨树县调查发现, 当地一次性施肥比例高达2/3以上^[6], 黄淮海玉米主产区亦如此 ...

2009

0.0

1.707

樊小林, 刘芳, 廖照源, 郑祥洲, 喻建刚. 我国控释肥料研究的现状和展望. 植物营养与肥料学报, 2009, 15: 463-473

Fan X

, Liu

, Liao Z

, Zheng X

, Yu J

The status and outlook for the study of controlled-release fertilizers in China.

Plant Nutr Fert Sci, 2009, 15: 463-473 (in Chinese with English abstract).

Based on years of experience on the study of controlled-release fertilizers and the extensive reference of both national and international research results, the research developments of controlled-release fertilizers in China were discussed by the authors. The academic value, economic significance, cutting-in and goal of the controlled-release fertilizer research in China were also analyzed and discussed. The R&D of controlled-release fertilizers with social demand and economic development were illustrated. The paper distinguished concept of controlled-release fertilizer from slow-release fertilizer, and discussed the significance to distinguish between controlled-release fertilizer and slow-release fertilizer. The history and status of controlled-release fertilizers research in China were reviewed. The history of research on the controlled-release fertilizers in China can be divided into 3 stages: the exploration period, the popularization period, and the integration and industrialization period for major technologies. Through the comparison of the controlled-release fertilizer research in China and abroad, the analysis on the trend and characteristics of the R&D of the controlled-release fertilizers in China and the discussion on the frontier field of controlled-release fertilizers, this article foresees the controlled-release fertilizer industry and indicates the direction for future research in China.

Fertilizer and Balanced Fertilization Institute, South China Agricultural University, Guangzhou 510642, China

本文结合作者长期从事控释肥料研究的心得和经验，在参阅国内外大量研究资料的基础上，比较系统地综述了我国控释肥料的研究进展。论文指出了我国控释肥料研究的学术价值和经济意义，阐述了我国控释肥料研发的切入点和目标、研制与社会需求、研制与我国经济发展。论述了控释肥料和缓释肥料的内涵和区别、区分控释肥料和缓释肥料的意义和作用。参阅1979年以来的研究资料，回顾了我国控释肥料研究的历史和现状，将我国控释肥料研制划分为3个阶段：1）缓控释肥料的探索阶段；2）控释肥料研究普及阶段；3）控释肥料关键技术深化和技术集成与产业化，概括了各个阶段的研究成果。在此基础上，通过对我国控释肥料研制与国际同类研究差距对比、我国控释肥料研究与发展总体趋势和特点分析、控释肥料研究的前沿领域之讨论，对我国控释肥料研究进行了展望，并提出了今后的研究方向。

... 如何协调肥料投入与产量之间的矛盾已引起广泛重视, 氮素释放速率与作物需肥规律基本一致、一次性基施满足生产需要、同时能有效提高生产效率、减少人工投入的缓控释肥的研发与应用被寄予厚望^{[7,8,9,10,11,12]} ...

2010

0.0

1.681

赵斌, 董树亭, 张吉旺, 刘鹏. 控释肥对夏玉米产量和氮素积累与分配的影响. 作物学报, 2010, 36: 1760-1768

Zhao

, Dong S

, Zhang J

, Liu

Effects of controlled- release fertilizer on yield and nitrogen accumulation and distribution in summer maize. Acta Agron Sin, 2010, 36: 1760-1768 (in Chinese with English abstract)

The field experiment was conducted to investigate the effects of controlled-release fertilizers (including one kind of fertilizer enveloped by colophony, CRF, and the other kind of fertilizer enveloped by sulfur, SCF) on accumulation and distribution of photosynthate and nitrogen as well as yield after anthesis of summer maize. Results indicated that photosynthetic rate of the treatments with controlled-release fertilizers CRF and SCF kept in a higher level after anthesis. Under the same application rates of N, P, and K, applying CRF (1 428 kg ha -1 ) and SCF (1 668 kg ha -1 ) increased the accumulation of dry matter and nitrogen per plant significantly compared with applying the common compound fertilizer (CCF, 1 260 kg ha -1 ). When the applied amounts of the CRF and SCF were decreased by 25%, the increment was also significant, and the distribution proportion of the dry matter from plant to seeds was significantly higher than that of applying CCF. The accumulation of the nitrogen in seeds of the CRF and SCF treatments was significantly higher than that of CCF treatment at maturity stage, and was increased with increasing applied proportion of fertilizer, but these was no significant difference between treatments of CRF, SCF and 25%.of CRF and SCF. Appling CRF and SCF increased the maize grain yield by 13.15% and 14.15% respectively under the same application rates. When the applied amount of CRF and SCF was decreased by 25%, the yield increment was 9.69% and 10.04%, respectively, the nitrogen use efficiency (NUE) and nitrogen agronomy efficiency were significantly higher than those of applying CCF.

State Key Laboratory of Crop Biology / College of Agronomy, Shandong Agricultural University, Tai’an 271018, China

在大田条件下研究树脂包膜控释肥(CRF)和硫包膜控释肥(SCF)对夏玉米花后光合产物和氮素积累与分配及产量的影响。结果表明，花后控释肥处理的光合速率始终较高，在相同施肥量(N、P、K量相同)情况下，控释肥CRF (1 428 kg hm -2 )和SCF (1 668 kg hm -2 )及其减量25%处理的单株干物质和氮素积累量都显著高于等量普通复合肥(CCF，1 260 kg hm -2 )处理；控释肥减量25%时，干物质向籽粒中的分配比例显著高于CCF处理；成熟期籽粒的氮素积累，控释肥处理都显著高于CCF处理，并随着控释肥比例的增加而增加，全量与减量25%的处理无显著差异。控释肥能显著提高玉米产量，等量控释肥增产13.15%和14.15%；控释肥减量25%时，分别比CCF增产9.69%和10.04%；其氮肥利用率和农学效率也均显著高于普通肥处理。

2013

3.534

0.0

... W3N3处理的产量没有继续增加, 表明其施氮量可能过量, 这与赵斌等^[9,11]和韩晓日等^[33]在控释尿素上的研究结果一致 ...

2001

0.0

1.707

符建荣. 控释氮肥对水稻的增产效应及提高肥料利用率的研究. 植物营养与肥料学报, 2001, 7: 145-152

Fu J

Effects of controlled release fertilizer on rice yield and N recovery.

Plant Nutr Fert Sci, 2001, 7: 145-152 (in Chinese with English abstract)

Field experiment was conducted at Tsuruoka City, Yamagata Prefecture, Japan in 1998 15 N trace method was used to test the effects of polyolefin coated urea (controlled release fertilizer) as basal on rice yield, the pattern of N absorption and N recovery rate compared with urea and ammonium sulfate with split application Results indicated that: 1) For transplanting rice, the highest yield was obtained by the blend of urea and LPS-100 in the ratio of 1:1 as basal, and increased by 23.6% and 9.2% compared with urea and ammonium sulfate, respectively For the direct seeding rice the highest yield was obtained by LP-100 as basal, and increased by 9.2% and 4.0% compared with urea and ammonium sulfate, respectively Controlled release fertilizer treatments could give more effective panicles and filled grains of rice than urea and ammonium sulfate treatments 2) No significant difference was observed in total N uptake among fertilizer treatments But the N accumulation from transplanting (or seeding) to panicle initiation in plants treated with controlled fertilizer were much more than conventional fertilization, resulting in the greater number of panicles per unit area and filled grains per panicle 3) 15 N labeled experiment indicated that up to maximum tiller stage N absorption of rice from basal fertilizer of urea and ammonium sulfate accounted for 33%～45% of total N absorption from basal fertilizer only N uptake from basal fertilizer lasted to heading stage in urea or ammonium sulfate plots and to harvesting stage in LP-100 plot While N uptake from topdressing applied at panicle initiation stage completed on 45 days after application 4) The total N recovery rate tested using 15 N method was higher in urea plot than ammonium sulfate and LP-100, while that tested using difference method was higher in the LP-100 than other treatments The N recovery rate of basal fertilizer was in the order of LP-100>ammonium sulfate> urea Basal application of urea/LPS-100 blend and LP-100 had higher physiological and agronomic efficiency of N than urea and ammonium sulfate, that may be the main reason for getting high yield by controlled fertilizer in the trail

Institute of soil and fertilizer;Zhejiang AAS;Hangzhou 310021;China

田间试验在日本山形县鹤冈市进行 ,探讨了树脂包膜类控释氮肥基施与尿素、硫铵分次施肥对单季稻的增产效应 ;应用 15 N示踪法研究了不同施肥处理水稻的吸氮模式及肥料利用率。结果表明 ,移栽稻以尿素与控释氮肥 (LPS-100)按 1∶1混合基施产量最高 ,分别比尿素和硫铵分次施肥增产23.6%和9.2% ;直播稻以LP-100基施产量最高 ,分别增产 9.2%和4.0% ;控释肥基施 ,水稻自移栽 (或直播 )至幼穗分化期吸氮量明显高于尿素或硫铵分次施肥。 15 N示踪研究表明 ,最高分蘖前水稻从尿素和硫铵基肥中吸收的氮素只占基肥氮总吸收量的 33%～ 45% ,水稻从基肥中吸收氮素可延续至抽穗期 ,而从LP-100中吸氮可持续至收获期。 15 N示踪法测得的氮素总回收率 ,以尿素处理最高 ,硫铵和LP-100接近 ;差减法测得的氮素总回收率以LP-100为最高 ,尿素 /LPS-100、尿素、硫铵三者接近 ;基肥的氮素回收率以LP-100>硫铵 >尿素。尿素 /LPS-100和LP-100一次基施 ,氮素生理效率和农学效率明显高于尿素和硫铵 ,是高产的主要原因

2009

0.0

1.904

... W3N3处理的产量没有继续增加, 表明其施氮量可能过量, 这与赵斌等^[9,11]和韩晓日等^[33]在控释尿素上的研究结果一致 ...

2001

0.0

2003

0.0

0.947

... 水氮供应是调控作物生长和发育的重要手段, 通过合理的水氮运筹调节作物光合特性及光合产物的积累和分配, 是实现作物高产的重要措施之一^[13] ...

2006

1.265

0.0

... 适时适度减少水分供应不会降低作物光合速率, 却显著减少作物蒸腾失水, 从而提高作物水分利用效率^[14] ...

1995

0.0

1.681

戴俊英, 沈秀瑛, 徐世昌, 王莲芝. 水分胁迫对玉米光合性能及产量的影响. 作物学报, 1995, 21: 356-363

Dai J

, Shen X

, Xu S

, Wang L

The effect of water stress on maize photosynthetic characters and yield.

Acta Agron Sin, 1995, 21: 356-363 (in Chinese with English abstract)

Shenyang Agricultural University, Shenyang,110161

... 徐世昌等研究表明水分胁迫导致玉米光合性能及产量的降低^[15] ...

2000

0.0

1.125

... 在节水节肥条件下, 通过氮水耦合也可以实现玉米高产^[16] ...

2002

0.0

1.681

李潮海, 刘奎, 周苏玫, 栾丽敏. 不同施肥条件下夏玉米光合对生理生态因子的响应. 作物学报, 2002, 28: 265-269

Li C

, Liu

, Zhou S

, Luan L

Response of photosynthesis to eco-physiological factors of summer maize on different fertilizer amounts.

Acta Agron Sin, 2002, 28: 265-269 (in Chinese with English abstract)

不同施肥水平下夏玉米光合对生理、生态因子的响应研究表明,施肥可以改善叶肉细胞的光合能力,使阻碍光合速率进一步提高的因素由非气孔限制逐渐转变为气孔限制,并可提高生育后期叶片的光合强度,延长高光合持续期,在日变化中表现为增加了叶片下午的光合强度.大田条件下,光合有效辐射(PAR)和气孔导度(S.)是影响光合速率(Pn)最主

... 李潮海等^[17]指出, 施肥能显著增加玉米的光合速率, 并且随着施肥量的增加后期高光合速率持续期延长 ...

... 常敬礼等研究表明氮肥提高了穗位叶叶片的净光合速率和叶绿素含量, 并且施氮肥越多, 各项光合参数值的增加幅度越大^[21,22,23], 李潮海等^[17]指出, 施肥可以改善叶肉细胞的光合能力, 提高生育后期叶片的光合强度, 延长高光合持续期 ...

1999

0.0

... 参考张守仁^[18]的方法计算叶绿素荧光动力学参数 ...

1994

0.0

1.125

... 1 控释尿素水氮耦合对夏玉米光合特性的影响作物产量的90%以上来自光合作用^[19], 光合作用是玉米干物质生产的基础, 其功能直接影响玉米籽粒产量 ...

1994

0.0

1.125

... 宋凤斌等研究认为, 水分胁迫下玉米叶片光合速率和气孔导度随胁迫增强而下降, 而在重度胁迫下光合速率显著下降, 水分胁迫使玉米叶片的叶绿素含量降低^[20] ...

2008

0.0

0.161

2012

0.0

1.904

王晓娟, 贾志宽, 梁连友, 丁瑞霞, 王敏, 李涵. 不同有机肥量对旱地玉米光合特性和产量的影响. 应用生态学报, 2012, 23: 419-425

Wang X

, Jia Z

, Liang L

, Ding R

, Wang

, Li

Effects of organic fertilizer application rate on leaf photosynthetic characteristics and grain yield of dry land maize.

Chin J Appl Ecol, 2012, 23: 419-425 (in Chinese with English abstract)

A 4-year field experiment was conducted at the Heyang Research Station in Weibei dryland to study the effects of organic fertilizer application rate on the leaf photosynthetic characteristics and grain yield of dryland maize. Comparing with applying chemical fertilizer, applying organic fertilizer increased the leaf photosynthetic rate and stomatal conductance, but decreased the leaf intercellular CO2 concentration at each growth stage of maize significantly. With the increasing application rate of organic fertilizer, the leaf photosynthetic rate and stomatal conductance at each growth stage of maize had a gradual increase, while the leaf intercellular CO2 concentration had a gradual decrease. The leaf photosynthesis of maize at each growth stage was controlled by non-stomatal factors, and the application of organic fertilizer reduced the non-stomatal limitation on the photosynthesis performance significantly. The 4-year application of organic fertilizer improved soil nutrient status, and soil nutrients were no longer the main factors limiting the leaf photosynthetic rate and grain yield of maize.

1Chinese Arid Area Research Institute of Water-Saving Agriculture, Northwest A & F University, Yangling 712100, Shaanxi, China; 2Ministry of Agriculture Key Laboratory of Crop Physi-ecology and Tillage Science in Northwestern Loess Plateau, Yangling 712100, Shaanxi, China; 3College of Resources and Environment, Northwest A & F University, Yangling 712100, Shaanxi, China

在渭北旱塬合阳试验基地进行了4年的旱地有机培肥试验, 研究不同有机肥施用量对旱地玉米光合特性和产量的影响.结果表明：与单施化肥相比，施有机肥处理的玉米各生育时期的叶片光合速率和气孔导度显著增大，胞间CO2浓度显著减小；随着有机肥施用量的增加，玉米叶片各生育时期的叶片光合速率和气孔导度逐渐增大，胞间CO2浓度逐渐减小.玉米各生育时期的光合作用主要受非气孔因素限制，施用有机肥显著降低了非气孔因素对光合的限制.连续4年施用有机肥，改善了土壤养分状况，使养分不再是玉米光合速率和产量的主要限制因子.

2007

0.0

1.289

段巍巍, 李慧玲, 肖凯, 李雁鸣. 氮肥对玉米穗位叶光合作用及其生理生化特性的影响. 华北农学报, 2007, 22(1): 26-29

Duan W

, Li H

, Xiao

, Li Y

Effects of nitrogen fertilizer on photosynthesis and its physiological and biochemical characteristics in ear-leaf of maize. Acta Agric Boreali-Sin, 2007, 22(1): 26-29 (in Chinese with English abstract)

The study was carried out in order to explore the responses of photosynthetic performances of summer maize to nitrogen nutrition. Four treatments were designed as 0, 112.5, 225 and 337.5 kg/ha respectively. The results showed that P(n), chlorophyll content and soluble protein increased with the addition of nitrogen fertilization, but the degradation of chlorophyll and soluble protein delayed. Hill reaction activity and PEPCase activity of nitrogen treatments were higher than that of CK for all periods, and the more the fertilizer input, the larger extent the photosynthetic parameters increased. The influence of fertilizer on P(n) was similar to those on Hill reaction activity and PEPCase activity on the whole, which suggested that fertilizer regulated P(n) by affecting electron transport capability and the photosynthetic carboxylase activity.

College of Agronomy, Agricultural University of Hebei, Baoding 071001, China

为明确氮素营养对玉米光合作用影响的可能机制，在河北农业大学标本园进行了田间试验。试验设4个施氮水平，分别为0，112.5，225和337.5 kg/hm 2 。对穗位叶光合作用参数的系统测定表明，在试验设计的施氮量范围内，氮肥提高了穗位叶叶片的Pn、叶绿素含量和可溶性蛋白质含量、延迟了叶绿素和可溶性蛋白质的分解。各测定时期，施用氮肥的处理，其叶片的Hill反应活力、PEPCase活性均高于对照，并且施氮肥越多，各项光合参数值的增加幅度越大，表观光合速率对氮肥的反应与各项光合生理生化指标的反应基本一致。说明氮肥通过调节光合电子传递能力和光合羧化酶的活性而影响表观光合速率的高低。

2009

0.0

1.289

齐华, 白向历, 孙世贤, 刘明, 杨国航, 丛雪. 水分胁迫对玉米叶绿素荧光特性的影响. 华北农学报, 2009, 24(3): 102-106

, Bai X

, Sun S

, Liu

, Yang G

, Cong

Effects of water stress on chlorophyll fluorescence parameters of maize.

Acta Agric Boreali-Sin, 2009, 24(3): 102-106 (in Chinese with English abstract)

Use maize Jidan 261 and Jingdan 28 as materials. Research the effects of water stress on chlorophyll fluO 2 rescence parameters of maize leaf. The result sho wed that the light2saturated net photosynthetic rate, light compensation point and light saturation point and values of intrinsic quantumefficiency for CO 2 uptake were decreased on water stress. The basis fluorescence of the maize leaveswas significantly increased. The max fluorescence, the variable fluorescence and photochemical efficiency were significantly decreased. With the increased of light, The photochemical quenching coeffi2 cient and actual photochemical efficiency were significantly decreased. But the non2photochemical quenching coefficient and apparent electron transport rate were increased slowly. Result showed that the photosynthesis competence ofmaize leaf was lo wer under stress.The inverse proportion of PS ò was unfolded slowly.The capability of electron transport of photO 2 synthesis was decreased, and PS ò latency active was restrained. The superabundance light was dissipation by other aP 2 proach such as non2photochemical quenching mode.To protected photosynthetic structures from injure.

1. College of Agronomy, ShenyangAgricultural University, Shenyang 110161, China; 2. National Agricultural Technology Extension and Service Center, Beijing 100026, China; 3. Maize Research Centre, Beijing Academy of Agriculture and Forestry Sciences, Beijing 100097, China

以吉单261和京单28两个玉米品种为试验材料,研究水分胁迫对玉米叶片光合特性及叶绿素荧光参数的影响.结果表明:水分胁迫下玉米叶片的最大净光合速率、光补偿点、光饱和点和表观量子效率均显著下降;基础荧光显著增加,最大荧光、可变荧光和PSⅡ光化学效率显著降低;随着光强增加,水分胁迫条件下的光化学猝灭系数和实际光化学效率下降幅度明显加剧;非光化学猝灭系数和表观电子传递速率增加幅度则明显减缓.表明水分胁迫使玉米叶片光合性能减弱,PsⅡ反应中心开放部分的比例减少,光合电子传递能力下降,PSⅡ潜在活性受到抑制,过多的光能多是以非光化学猝灭等其他形式耗散掉,从而保护光合机构免受伤害.

... 2 控释尿素水氮耦合对夏玉米荧光特性的影响通过叶绿素荧光可快速检测完整植株在水分胁迫下光合作用的真实行为, 用来评价光合机构的功能和环境胁迫的影响^[24] ...

... 齐华等^[24]研究表明水分胁迫使玉米叶片光合性能减弱, PSII反应中心开放部分的比例减少, 光合电子传递能力下降, PSII潜在活性受到抑制, F_v/F_m、F_m和F_v降低, F_o升高, q_P降低, NPQ升高, 王建程等^[25]研究发现提高施肥水平有利于减轻水分胁迫对光合中心的伤害, 高肥有助于提高玉米PSII光化学效率 ...

2005

0.0

1.835

2009

0.0

1.681

李耕, 高辉远, 赵斌, 董树亭, 张吉旺, 杨吉顺, 王敬锋, 刘鹏. 灌浆期干旱胁迫对玉米叶片光系统活性的影响. 作物学报, 2009, 35: 1916-1922

, Gao H

, Zhao

, Dong S

, Zhang J

, Yang J

, Wang J

, Liu

Effects of drought stress on activity of photo systems in leaves of maize at grain filling stage.

Acta Agron Sin, 2009, 35: 1916-1922 (in Chinese with English abstract)

At grain filling stage, the effects of drought stress on photosynthetic acivities of photosystem I (PS I) and photosystem II (PS II) in leaves of maize (Zhengdan 21, a cultivar with high starch content) were studied by simultaneously analyzing chlorophyll a fluorescence transient and light absorbance at 820 nm. The results, obtained from two years experiments, demonstrated that the drought stress significantly reduced photosynthesis ( P P A in PS II. The acceptor side of PSII was damaged more severely than the donor side of PSII. Furthermore, the maximal oxidation-reduction activity of PS I ( Δ I/I o ) was also significantly decreased by the drought stress, which inhibited the photosynthetic electron tranport from the PS II to PS I, destructing the coordination between PS I and PS II. We suggest that the inhibition of PS I and PS II and the destruction of the coordination between PS I and PS II by the drought stress is one of the main reasons to cause the decrease in photosyntheis and grain yield of maize.

1 College of Agronomy, Shandong Agricultural University; 2 College of Life Science, Shandong Agricultural University; 3 State Key Laboratory of Crop Biology, Tai’an 271018, China

以高淀粉玉米品种郑单 21 为材料，借助叶绿素荧光快速诱导动力学曲线和 820 nm 光吸收曲线，研究了灌浆期土壤干旱胁迫对玉米籽粒产量和对叶片光系统 I (PSI) 及光系统 II (PS II) 活性的影响。两年的研究结果均表明，干旱胁迫显著抑制叶片光合速率 ( P 和籽粒产量 ( P 。 JIP-test 分析发现干旱胁迫导致叶绿素荧光快速诱导动力学曲线中的 K 点和 J 点上升，表明 PS II 放氧复合体 (OEC) 和 Q A 之后的电子传递链受到抑制，且 PS II 受体侧受抑制的程度大于供体侧。此外，干旱胁迫也显著地抑制 PS I 的最大氧化还原活性 ( Δ I/I o ) ，阻碍光合电子从 PS II 向 PS I 的传递，破坏了 PS I 和 PS II 的协调性。我们认为干旱胁迫抑制 PS I 和 PS II 活性并破坏二者的协调性，是导致导致 P n 和籽粒产量下降的重要原因之一。

... W1, 对水分效应敏感, 这与李耕等^[26]运用JIP-test分析研究干旱胁迫对玉米叶片光系统活性影响的结果一致 ...

2001

0.0

1.125