<graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="0496-3490-40-731/img_1.png"/>不同土层测墒补灌对小麦旗叶光合特性和干物质积累与分配的影响

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郭增江, 于振文, 石玉, 赵俊晔, 张永丽, 王东. 不同土层测墒补灌对小麦旗叶光合特性和干物质积累与分配的影响 . 作物学报, 40(4): 731-738
GUO Zeng-Jiang, YU Zhen-Wen, SHI Yu, ZHAO Jun-Ye, ZHANG Yong-Li, WANG Dong. Photosynthesis Characteristics of Flag Leaf and Dry Matter Accumulation and Allocation in Winter Wheat under Supplemental Irrigation after Measuring Moisture Content in Different Soil Layers. 作物学报, 40(4): 731-738 复制到剪切板

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不同土层测墒补灌对小麦旗叶光合特性和干物质积累与分配的影响

郭增江¹, 于振文¹, 石玉^1,^*, 赵俊晔², 张永丽¹, 王东¹

¹山东农业大学 / 农业部作物生理生态与耕作重点实验室, 山东泰安271018

² 中国农业科学院农业信息研究所, 北京 100081

^* 通讯作者(Corresponding author): 石玉, E-mail:shiyu@sdau.edu.cn, Tel: 0538-8241484

收稿日期:2014-01-12

基金:本研究由国家自然科学基金项目(31171498)和山东省自然科学基金项目(ZR2011CQ014)资助。

摘要

2011—2012和2012—2013年连续2个小麦生长季, 在大田条件下, 设置0~20 cm (D1)、0~40 cm (D2)、0~60 cm (D3)和0~140 cm (D4) 4个土层测定土壤含水量, 以各土层平均土壤相对含水量拔节期65%和开花期70%为目标相对含水量, 全生育期不灌溉为对照处理 (D0), 研究依据不同土层的土壤含水量测墒补灌对小麦旗叶光合特性和干物质积累与分配的影响。结果表明: D2的开花期叶面积指数和单位土地面积上旗叶叶面积、开花后7 d和14 d的旗叶净光合速率和实际光化学效率均高于其他处理, 而气孔限制值低于其他处理; D2的成熟期干物质积累量、开花后干物质向籽粒的分配量和开花后同化物分配对籽粒的贡献率亦高于其他处理。两年度D2的籽粒产量分别为9367.4 kg hm^-2和9727.5 kg hm^-2, 均显著高于其他处理; 同时, D2的水分利用效率高于D0、D3和D4处理, 与D1处理无显著差异。因此, 于小麦拔节期和开花期依据0~40 cm土层的土壤含水量测墒补灌是同步实现高产和高水分利用效率的有效措施。

关键词: 小麦; 测墒补灌; 光合特性; 干物质积累与分配; 籽粒产量

Photosynthesis Characteristics of Flag Leaf and Dry Matter Accumulation and Allocation in Winter Wheat under Supplemental Irrigation after Measuring Moisture Content in Different Soil Layers

GUO Zeng-Jiang¹, YU Zhen-Wen¹, SHI Yu^1,^*, ZHAO Jun-Ye², ZHANG Yong-Li¹, WANG Dong¹

¹ Shandong Agricultural University / Key Laboratory of Crop Ecophysiology and Farming System, Ministry of Agriculture, Tai#cod#x02019;an 271018, China

² Institute of Agricultural Information, Chinese Academy of Agricultural Sciences, Beijing 100081, China

Fund:

Abstract

This study aimed to propose a suitable soil layer depth used in determining the irrigation amount. A field experiment was conducted in the 2011-2012 and 2012-2013 wheat growing seasons to study the effects of supplemental irrigation based on the measurement of soil moisture contents on photosynthesis of flag leaf and dry matter accumulation and allocation. Four irrigation treatments were designed with target soil moisture of 65% at jointing and 70% at anthesis in 0-20 (D1), 0-40 (D2), 0-60 (D3), and 0-140 cm (D4) soil layers. Zero-irrigation (D0) was used as the control. D2 was superior to other treatments with higher values of leaf area index (LAI) and flag leaf area on one square meter land at anthesis, photosynthetic rate (P_n) and actual photochemical efficiency (Φ_PSII) at seven and fourteen days after anthesis; whereas, the stomatal limitation (L_s) of D2 was lower than those of other treatments. Compared with other treatments, D2 had larger dry matter accumulation at maturity, more dry matter allocated in grains, and higher contribution ratio of dry matter from vegetative organs to grain after anthesis. The grain yield in D2 was 9367.4 kg ha^-1 in 2011-2012 growing season and 9727.5 kg ha^-1 in 2012-2013 growing season, which were significantly higher than those in other treatments. The water use efficiency of D2 was significantly higher than those of D0, D3, and D4, but with out significant difference to that of D1. To obtain both high yield and high water use efficiency, we suggest the optimal soil layer for measuring moisture content is 0-40 cm, and supplementary water should be given at jointing and anthesis based on measured soil moisture.

Keyword: Winter wheat; Supplemental irrigation based on measurement of moisture content; Photosynthesis characteristics; Dry matter accumulation and allocation; Grain yield

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黄淮海地区人均水资源占有量为519 m³, 仅为全国的1/5^{[ 1, 2]}, 小麦生长季降水量不超过200 mm, 水分蒸散量约为400~500 mm^{[ 3]}, 水资源短缺成为制约该地区小麦生产的主要限制因子。水分亏缺显著降低旗叶净光合速率^{[ 4]}, 但适度水分亏缺可以提高作物产量和水分利用效率^{[ 5]}。试验表明, 拔节期和开花期是通过灌水以提高产量及水分利用效率的关键时期^{[ 6]}。仅拔节期灌水75 mm, 小麦旗叶气孔导度下降, 蒸腾速率随之降低, 与拔节期、开花期各灌溉75 mm处理相比, 净光合速率低5.3%^{[ 7]}。小麦拔节期和开花期分别灌溉60 mm, 开花后净光合速率显著提高, 较仅拔节期灌溉60 mm的处理高16.6%^{[ 8]}。亦有研究指出, 拔节期、孕穗期和灌浆期各灌一次水, 每次60 mm, 小麦灌浆中期的净光合速率高于拔节期和孕穗期灌水, 其中净光合速率升高约10 μmol m^-2 s^{-1 [ 9]}。小麦拔节期与开花期各灌溉60 mm使花前干物质转运量较仅拔节期灌溉每株减少1.3 g, 但成熟期干物质积累量显著高于拔节期灌溉处理, 每株增加2.4 g^{[ 10]}。在小麦越冬期和拔节期各灌水75 mm条件下, 开花后干物质同化量较越冬期灌溉 75 mm的处理提高19.8%, 产量增加2068.7 kg hm^{-2 [ 11]}。前人研究小麦旗叶光合特性及干物质积累与分配多采用定量灌溉, 本试验依据不同土层的土壤含水量进行测墒补灌, 以期为小麦节水高产栽培提供理论依据。

1 材料与方法

1.1 试验设计

2011—2012和2012—2013年冬小麦生长季, 在山东兖州小孟镇史王村大田(35°40'09''N, 116°41'43''E)进行田间试验, 试验田土壤质地为壤土。2011—2012年度播种前试验田0~20 cm土层含有机质1.39%、全氮0.12%、碱解氮142.29 mg kg^-1、速效磷31.01 mg kg^-1和速效钾 112.60 mg kg^-1。小麦生育期间降雨量为183 mm, 其中播种至拔节期152 mm, 拔节至开花期31 mm, 开花至成熟期0 mm。2012—2013年度播种前试验田0~20 cm土层含有机质1.35%、全氮0.12%、碱解氮143.63 mg kg^-1、速效磷34.51 mg kg^-1和速效钾118.48 mg kg^-1。小麦生育期间降雨量为238 mm, 其中播种至拔节期92 mm, 拔节至开花期32.5 mm, 开花至成熟期113.5 mm。0~140 cm各土层的田间持水量和容重见表1。

小麦品种为济麦22。播种前底施纯氮105 kg hm^-2、P₂O₅和K₂O各150 kg hm^-2, 拔节期开沟追施纯氮 135 kg hm^-2; 肥料为尿素、磷酸二铵和氯化钾。分别于2011年10月12日和2012年10月9日播种, 2012年6月12日和2013年6月14日收获。三叶期定苗, 留苗密度为180株 m^-2。其他管理措施同一般高产田。小区面积4 m × 4 m = 16 m², 随机区组排列, 3次重复, 小区之间留1.5 m宽隔离区, 并播种小麦。

设不灌溉对照(D0)和4个测墒土层处理, 分别是0~ 20 cm (D1)、0~40 cm (D2)、0~60 cm (D3)和0~140 cm (D4)。以平均土壤相对含水量拔节期65%和开花期70%为土壤湿度目标, 于拔节期和开花期灌水前测墒, 计算灌水量, 并于灌水后3 d取土测定实际土壤含水量。灌水量计算公式为 M = 10 γH( β_i - β_j)^{[ 12]}, 式中, M为灌水量(mm), H为土壤计划湿润层的深度(cm), γ为计划湿润层的土壤容重(g cm^-3), β_i为目标含水量(田间持水量乘以目标相对含水量), β_j为灌溉前土壤含水量。用水表计量实际灌水量。

2011—2012年度, 测墒补灌测得的土壤含水量与设计目标的相对误差(以下称“调控误差”)平均为2.0%, 2012—2013年度, 各处理的平均调控误差为2.3% (表1), 表明本试验中采取的测墒补灌方法能够实现预期设计。

1.2 测定项目与方法

1.2.1 土壤相对含水量的计算用土钻取0~200 cm土层的土壤, 每20 cm为一层土样, 取土后立即装入铝盒, 称鲜重, 105℃烘干至恒重, 称干重, 计算土壤含水量和土壤相对含水量。土壤含水量(%) = (土壤鲜重 - 土壤干重) / 土壤干重 × 100; 土壤相对含水量 (%) = 土壤含水量 / 田间持水量 × 100。

1.2.2 农田耗水量和水分利用效率的计算根据水分平衡法计算小麦生育期总耗水量, ET=Δ S+ M+ P+ K^{[ 13, 14]}。式中, ET为总耗水量(mm); Δ S为小麦生育期间土壤贮水的变化量(mm); M为灌水量(mm); P为降水量(mm); K为地下水补给量(mm), 当地下水埋深大于2.5 m时, K值可以不计。本试验地区地下水埋深在4 m以下^{[ 15]}, 因此无地下水补给。

水分利用效率 = 籽粒产量 / 作物全生育期耗水量^{[ 16, 17]}。

1.2.3 叶面积指数和单位土地面积上旗叶叶面积测定

依据Li等的方法^{[ 18]}, 叶面积 = 最大叶长×最大叶宽×0.78, A_{f, PA}= D × A_f, 式中, A_{f, PA}为单位土地面积上旗叶叶面积(m² m^-2), D为群体密度(单茎m^-2), A_f为单茎旗叶叶面积(m²)。叶面积指数(LAI)为单位土地面积上所有绿叶面积之和与占土地面积的比值。

1.2.4 旗叶净光合速率、气孔限制值和实际光化学效率测定

用CIRAS-2型光合作用测定系统(英国产)和FMS-2型荧光仪于开花后7 d和14 d的9:00-11:00, 自然光照下测定旗叶净光合速率( P_n)、胞间二氧化碳浓度( C_i)和大气中二氧化碳浓度( C_a)及旗叶实际光化学效率( Φ_PSII)。气孔限制值 L_s= 1- C_i/ C_a^{[ 19]}。

1.2.5 干物质测定于开花期和成熟期取样, 成熟期按叶片、茎秆+叶鞘、颖壳+穗轴、籽粒等器官取样, 并在70℃条件下, 烘干至恒重, 称干物质质量。各指标计算公式^{[ 8]}如下。

开花前营养器官贮藏同化物转运量=开花期干物质量-成熟期干物质量;开花前营养器官贮藏同化物对籽粒贡献率(%)=(开花期干重-成熟期干重)/成熟期籽粒干重×100;花后同化物在籽粒中的分配量=成熟期籽粒干重-开花前营养器官贮藏同化物转运量;花后同化物对籽粒贡献率(%)=花后同化物在籽粒中的分配量/成熟期籽粒干重×100。

1.2.6 籽粒产量于小麦成熟期小区收获、脱粒、自然风干后测产, 籽粒水分含量为12.5%。

表1 不同处理的目标相对含水量、灌溉后实际相对含水量和灌溉量 Table 1 Target relative water content, relative soil water content after irrigation, and irrigation amounts in the different treatments

1.3 数据处理

用Microsoft Excel 2003和SigmaPlot 10.0软件计算数据和绘图, 用DPS 7.05统计软件进行显著性检验(LSD法)。

2 结果与分析

2.1 不同处理对开花期叶面积指数和单位土地面积上旗叶叶面积的影响

2012—2013年度开花期叶面积指数, D2显著高于D0、D1、D3和D4处理, D1和D3间无显著差异, 均高于D0和D4处理, D4高于D0处理; 单位土地面积上旗叶叶面积, D2高于其他处理, D1和D3间无显著差异, 高于D4和D0处理, D0处理最低。2011—2012年度结果与此一致(图1)。说明依据0~40 cm土层的土壤含水量测墒补灌, 提高了小麦开花期叶面积指数和单位土地面积上的旗叶叶面积。

2.2 不同处理对小麦旗叶 P_n、 L_s和 Φ_PSII的影响

开花后7 d和14 d的旗叶 P_n, D2显著高于其他处理, D1和D3高于D4处理, D0处理最低(图2-A, D)。花后7 d的旗叶 L_s表现为D0、D4>D1、D3>D2, D0与D4间、D1与D3间无显著差异(图2-B); 花后14 d的 L_s, D0高于其他处理, D1、D3和D4间无显著差异, 高于D2处理(图2-E)。花后7 d和14 d的旗叶 Φ_PSII, D2高于其他处理, D1与D3间无显著差异, 高于D0和D4处理, D0处理最低(图2-C, F)。表明依据0~40 cm土层的土壤含水量测墒补灌, 提高了小麦旗叶光合能力, 有利于碳水化合物的合成。

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	图1 不同处理开花期的叶面积指数(A)和单位土地面积上旗叶叶面积(B)(2012-2013)Fig. 1 Leaf area index (A) and flag leaf area on one square meter land (B) at anthesis in wheat with different treatments (2012-2013)

误差线上不同字母表处理间差异显著(LSD, P<0.05)。

Different letters above error bars indicate significant difference among treatments by LSD test ( P<0.05).

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	图2 不同处理小麦开花后7 d (A、B、C)和14 d (D、E、F)的旗叶 P_n、 L_s和 Φ_PSII (2012-2013)Fig. 2 Net photosynthetic rate ( P_n), stomatal limitation ( L_s), and actual photochemical efficiency ( Φ_PSII) of flag leaf at seven days (A, B, C) and fourteen days (D, E, F) after anthesis in wheat with different treatments (2012-2013)

误差线上不同字母表处理间差异显著(LSD, P<0.05)。

Different letters above error bars indicate significant difference among treatments by LSD test ( P<0.05).

2.3 不同处理对小麦开花期和成熟期干物质积累的影响

2011—2012年度开花期干物质积累量, D2与D3间无显著差异, 高于D0、D1和D4处理, D1高于D0和D4处理, D0处理最低; 成熟期各处理的干物质积累量为D2>D1、D3>D4>D0, D1与D3间无显著差异。2012—2013年度的开花期干物质积累量, D2高于其他处理, D1和D4间无显著差异, 低于D3处理, D0处理最低; 成熟期干物质积累量, D2高于其他处理, D1与D3间无显著差异, 高于D0和D4处理, D4高于D0处理(图3)。依据0~40 cm土层的土壤含水量测墒补灌, 促进了干物质的合成, 利于成熟期干物质的积累。

2.4 不同处理对小麦成熟期干物质在各器官中分配量的影响

各处理干物质在籽粒和茎秆+叶鞘中的分配量均表现为D2>D3>D1>D4>D0; D2的干物质在颖壳+穗轴中的分配量高于其他处理, D1和D3间无显著差异, 高于D0和D4处理, D0处理最低; 干物质在叶片中的分配量为D2>D1、D3>D0、D4, D1与D3间、D0与D4间无显著差异。两年度结果一致(图4)。依据0~40 cm土层的土壤含水量测墒补灌, 各器官中同化物的分配量提高, 并有利于同化物向籽粒中转运及分配, 提高籽粒产量。

2.5 不同处理对开花后营养器官干物质再分配量的影响

2011—2012年度, 各处理花前营养器官贮藏同化物转运量为D0>D1>D3、D4>D2, D1与D3间无显著差异; 花前营养器官贮藏同化物对籽粒贡献率以D0高于其他处理, D1和D4高于D2和D3处理。各处理花后贮藏同化物在籽粒中的分配量和对籽粒贡献率分别为D2>D3>D1>D4> D0和D2>D3>D1、D4>D0 (表2)。

2012—2013年度花前营养器官贮藏同化物转运量, D0高于其他处理, D1次之, D2最低; 各处理花前营养器官贮藏同化物对籽粒贡献率为D0>D1、D4>D3>D2, D1与D4间无显著差异。花后干物质在籽粒中的分配量为D2> D3>D1>D4>D0; 花后干物质分配量对籽粒的贡献率为D2>D3>D1、D4>D0。表明依据0~40 cm土层的土壤含水量测墒补灌, 促进了开花后同化物的积累及其向籽粒中的分配, 是籽粒产量提高的生理基础。

2.6 不同处理对籽粒产量、总耗水量和水分利用效率的影响

两年度籽粒产量均为D2>D3>D1>D4>D0 (表3)。2011—2012年度的耗水量, D2高于其他处理, D1和D4低于D3处理, D0处理最低。水分利用效率, D1和D2间无显著差异, 高于D0、D3和D4处理, D3高于D0和D4处理。2012—2013年度的耗水量, D3高于其他处理, D2高于D0、D1和D4处理, D1与D4高于D0处理。水分利用效率为D1、D2>D3>D0、D4, D1与D2间、D0与D4间无显著差异。因此测墒土层深度以0~40 cm最佳, 可以获得高产和高水分利用效率。

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	图3 不同处理小麦开花期和成熟期的干物质积累量误差线上不同字母表处理间差异显著(LSD, P<0.05)。Fig. 3 Dry matter accumulation amounts of wheat with different treatments at anthesis and maturityDifferent letters above error bars indicate significant difference among treatments by LSD test ( P<0.05).

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	图4 不同处理成熟期干物质在各器官中的分配量(2011-2012)Fig. 4 Dry matter allocation amounts in different organs at maturity with different treatments (2011-2012)

误差线上不同字母表处理间差异显著(LSD, P<0.05)。

Different letters above error bars indicate significant difference among treatments by LSD test ( P<0.05).

3 讨论

灌溉显著影响开花后小麦旗叶的光合能力, 拔节期和开花期各灌溉75 mm的开花后净光合速率较仅拔节期灌溉75 mm的处理高4.9 μmol m^-2 s^{-1 [ 7]}。开花后水分亏缺, 旗叶光合速率和气孔导度降低, 气孔限制值升高, 导致实际光化学效率从0.8降低至0.4^{[ 20]}。在拔节期和开花期分别灌溉60 mm, 开花后小麦叶片的光合速率提高, 光合功能持续期延长, 光合速率为13.53 μmol m^-2 s^-1, 比仅拔节期灌溉60 mm的处理提高14.3%^{[ 8]}。不同供水条件下, 随灌水量减少各器官气孔密度增大, 其中旗叶增幅较大, 约9%, 与气孔导度呈负相关^{[ 21]}, 导致气孔限制值升

表2 不同处理开花后营养器官干物质的再分配量 Table 2 Dry matter allocation amounts from vegetative organs after anthesis with different treatments

表3 不同处理的籽粒产量、总耗水量和水分利用效率 Table 3 Grain yield, evapotranspiration, and water use efficiency of winter wheat with different treatments

高; 与固定根区灌溉处理相比, 交替根区灌溉的叶片净光合速率升高5 μmol m^-2 s^{-1 [ 22]}。Zhao等^{[ 23]}研究指出, 拔节期和抽穗期分别灌溉60 mm, 小麦抽穗期叶面积指数显著提高, 达到8.26; 拔节期灌溉75 mm, 小麦孕穗期和开花期叶面积指数分别为6.8和5.2^{[ 10]}。本试验条件下, 依据0~20 (D1)、0~40 (D2)、0~60 (D3)和0~140 (D4) cm土层的土壤含水量测墒补灌, 拔节期灌溉量分别为33.0、43.6、65.6和24.5 mm, 开花期D2处理的小麦叶面积指数和单位土地面积上旗叶叶面积均高于其他处理, 开花后气孔限制降低, 旗叶净光合速率和实际光化学效率升高, 说明D2处理利于开花后光合作用, 促进碳水化合物的合成。

与抽穗期和灌浆期各灌溉60 mm的处理相比, 拔节期和抽穗期各灌溉60 mm的开花后单株地上部干物质积累量增加8.1 g, 干物质转移量和转移效率分别提高3.5 g和9.1%^{[ 24]}。与水分充足处理相比, 小麦开花后水分亏缺, 导致开花后16 d籽粒中同化物降低22%, 开花后32 d降低48%^{[ 25]}。灌浆前期, 籽粒中同化物占总生物量的35.25%~45.66%, 颖壳与穗轴占7.40%~9.85%和茎秆占27.58%~39.75%; 灌浆后期, 70%以上同化物运送至籽粒中^{[ 26]}。研究表明指出, 拔节期灌溉60 mm小麦单株生物量可达35.16 g, 且花前干物质积累及向籽粒再转运量较大, 对籽粒灌浆贡献率为47%^{[ 8]}; 亦有研究表明, 花前累积干物质对小麦产量有一定贡献, 但产量高低主要取决于花后干物质的积累和转移, 花后转移干物质贡献率可达65%以上^{[ 27]}。本研究依据0~40 cm土层的土壤含水量测墒补灌, 成熟期干物质的积累量和籽粒中干物质分配量高于其他处理, 花后营养器官贮藏同化物向籽粒的转运量及对籽粒的贡献率最高, 是获得最高籽粒产量的生理基础。

随着灌水次数的增加, 灌溉量从80 mm升高至 240 mm, 小麦耗水量增加80~90 mm, 水分利用效率降低0.3~0.4 kg m^{-3 [ 28]}。在底墒充足的条件下, 在拔节期和开花期分别灌水60 mm, 籽粒产量显著提高, 为7181 kg hm^-2, 水分利用效率为1.42 kg m^{-3 [ 29]}。Li等^{[ 6]}指出, 拔节期和抽穗期各灌溉60 mm小麦籽粒产量显著高于仅拔节期灌溉处理, 产量增加70.0 g m^-2, 水分利用效率亦有所升高。在本研究中, 两年度D2处理的籽粒产量最高, 分别为9367.4 kg hm^-2和9727.5 kg hm^-2, 同时获得高的水分利用效率。因此, 0~40 cm土层是测墒补灌的最适宜测定土层。本研究设定的目标土壤相对含水量为拔节期65%、开花期70%, 仅适于壤土条件, 而在沙土或黏土质地上如何应用测墒补灌技术, 有待进一步试验。

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.

参考文献

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2006

0.0

... 黄淮海地区人均水资源占有量为519 m³, 仅为全国的1/5 ^[1,2], 小麦生长季降水量不超过200 mm, 水分蒸散量约为400~500 mm ^[3], 水资源短缺成为制约该地区小麦生产的主要限制因子 ...

2006

0.0

2012

0.0

2008

0.0

1.667

. 2008, 34:1213-1219

Changes of chloroplast ultramicrostructure and function of different green organs in wheat under limited irrigation

(不同水分供给对小麦叶与非叶器官叶绿体结构和功能的影响) . (作物学报) , 2008 , 34 : 1213 - 1219 Zhang Y P , Wang Z M , Huang Q , Xie M . Changes of chloroplast ultramicrostructure and function of different green organs in wheat under limited irrigation

Zhang Y P

(张永平) , (王志敏) , (黄琴) , (谢岷) . (不同水分供给对小麦叶与非叶器官叶绿体结构和功能的影响) . (作物学报) , 2008 , 34 : 1213 - 1219 Zhang Y P

1 College of Agronomy and Bio-technology, China Agricultural University, Beijing 100094; 2 College of Agronomy, Inner Mongolia Agricultural University, Huhhot 010018, Inner Mongolia, China

The green organs in wheat (Triticum aestivum L.) consist of leaf and non-leaf organs including ear, culm, and sheath. Understanding the structures and functions of these non-leaf green organs under abiotic stresses will help to explore the potential of photosynthesis of the whole plant. In order to identify the photosynthetic adaptation of different green organs to drought stress in winter wheat, the changes of chloroplast ultra-microstructure and function of the organ above were studied in a field experiment in the 2002–2003 growth season (a medium rain fall growth season) using cultivar Shijiajuang 8 with water treatments of no irri-gation (control), irrigating twice at jointing and anthesis stages, and irrigating 4 times at pseudostem erecting, booting, anthesis, and grain-filling stages. Each irrigation received 750 m³ ha^-1 water. Chloroplast ultramicrostructure of flag leaf blade and non-leaf organs (peduncle, flag leaf sheath, glume, and lemma) in response to different irrigation treatments were observed during late grain-filling period, and the net photosynthetic rate (Pn) and Fv/Fm values of different green organs were measured simultaneously. The results showed that the chloroplast number of non-leaf organs was less than that of leaf blade, and many starch grains were found in the chloroplasts of glume and lemma under sufficient water supplies (irrigating 4 times). Obvious senescence characteris-tics and differences in chloroplast ultramicrostructure were found in different organs of control, and the senescence was signifi-cantly severer in flag leaf than in non-leaf green organs. The chloroplast structure of flag leaf blade and lemma was destroyed under drought stress, while that of peduncle, sheath, and glume showed relatively stable in the late developmental stage under different irrigation regimes. Post-anthesis drought stress decreased P_n and F_v/F_m values in different green organs, but much less affected them in non-leaf organs (peduncle, sheath, and ear). The results indicated greater stability of chloroplast structure and photosynthetic activity in non-leaf green organs at the late grain-filling stage under water stress.

在田间不同灌水条件(春季不灌水、春季灌2水和春季灌4水, 每次灌水750 m3 hm^-2)下, 于灌浆后期对小麦叶片与非叶器官(穗颖、穗下节和叶鞘)叶绿体结构进行了电镜观察, 并对不同器官的净光合速率(P_n)和光化学效率(F_v/F_m)进行了测定, 以期探明小麦叶与非叶器官光合结构和功能对水分变化的适应性差异。结果表明, 在充足供水条件(春季灌4水)下, 叶片的叶绿体数目明显多于各非叶器官, 但护颖和外稃叶绿体含有较多的淀粉粒。在灌浆期上层土壤重度水分亏缺(春季不灌水)条件下, 植株各器官叶绿体均出现明显的衰老特征, 但衰老程度存在器官间显著差异, 旗叶叶片叶绿体受损程度明显大于非叶器官。在所测非叶器官中, 外稃叶绿体对干旱胁迫反应较为敏感, 而护颖、穗下节间和旗叶鞘叶绿体结构具有较强的稳定性。水分胁迫下各器官P_n和F_v/F_m值均降低, 叶片降低幅度最大, 且随灌浆进程其光合下降最快, 但穗下节间、旗叶鞘和穗器官较为稳定, 可能与其叶绿体结构的稳定性有关。说明小麦非叶绿色器官光合结构与功能对水分亏缺具有较强的耐性。

... 水分亏缺显著降低旗叶净光合速率^[4], 但适度水分亏缺可以提高作物产量和水分利用效率^[5] ...

2003

0.0

. 2003, 23:2342-2352

The effect of limited water supply on root growth and soil water use of winter wheat

(人工控制有限供水对冬小麦根系生长及土壤水分利用的影响) . (生态学报) , 2003 , 23 : 2342 - 2352 Liu G S , Guo A H , Ren S X , An S Q , Lin R N , Zhao H R . The effect of limited water supply on root growth and soil water use of winter wheat

Liu G S

(刘庚山) , (郭安红) , (任三学) , (安顺清) , (林日媛) , (赵华荣) . (人工控制有限供水对冬小麦根系生长及土壤水分利用的影响) . (生态学报) , 2003 , 23 : 2342 - 2352 Liu G S

试验在中国气象局固城农业气象试验基地的大型人工控制农田水分试验场进行.在底墒充足的条件下采用3种供水处理:拔节期一次性供水75mm(I1);返青供水37.5mm和拔节期供水37.5mm(I2);返青后生长期内无水分供给(ICK).全生育期内用电动防雨棚遮去自然降水.试验结果表明,表层土壤(0～30cm)水多根多,根系反应敏感.上层干旱促使根系向深层发育,利用下层水量较多.I1处理减少表土层(0～30cm)的根量和根长密度,促进根系下扎,较多地利用深层土壤水分,并减少无效分蘖.虽然总穗数减少,但同I2、ICK相比籽粒数和籽粒重有较大幅度增加,提高了产量和水分利用率.根系吸水效率随土层深度增加呈下降趋势,I1在30cm以下其根系吸水效率超过了I2处理,并在100～200cm土层表现最为明显.ICK除0～30cm土层外,其余土层有效底墒供水率均较低;I1和I2两处理30～100cm有效底墒供水率均在84%以上,1～2m土层内I1大于I2.ICK由于土壤水分不足并未造成千粒重的明显下降,相反,3个处理中最高;土壤水分不足导致穗数、穗粒重、籽粒重和籽粒数的显著降低,从而造成了最终产量的降低.产量水平上的水分利用效率I1比其余处理提高了将近14%,其次为ICK,最低的是I2.在灌溉效率上,I1比I2提高了将近19%;在相同灌溉量的前提下,蒸腾效率I1比I2提高约7%.I1水分利用效率高于I2和I CK处理.可以认为:配合充足底墒,前期控水,有限的一定量的水分拔节期(关键期)一次供给比在返青和拔节分别供给,更为有效科学.

... 水分亏缺显著降低旗叶净光合速率^[4], 但适度水分亏缺可以提高作物产量和水分利用效率^[5] ...

2010

0.0

... 试验表明, 拔节期和开花期是通过灌水以提高产量及水分利用效率的关键时期^[6] ...

... Li等^[6]指出, 拔节期和抽穗期各灌溉60 mm小麦籽粒产量显著高于仅拔节期灌溉处理, 产量增加70 ...

2010

0.0

1.883

. 2010, 16:852-858

Effects of water-saving irrigation on water consumption and photosynthetic characteristics of winter wheat in Huang-Huai-Hai area of China

(节水灌溉对黄淮海地区冬小麦水分消耗与光合特性的影响) . (植物营养与肥料学报) , 2010 , 16 : 852 - 858 Ning D F , Li Z J , Sun W Y , Ma W P , Huang S W , Zhao B Q . Effects of water-saving irrigation on water consumption and photosynthetic characteristics of winter wheat in Huang-Huai-Hai area of China

Ning D F

(宁东峰) , (李志杰) , (孙文彦) , (马卫萍) , (黄绍文) , (赵秉强) . (节水灌溉对黄淮海地区冬小麦水分消耗与光合特性的影响) . (植物营养与肥料学报) , 2010 , 16 : 852 - 858 Ning D F

2008～2009年在大田试验条件下研究了节水灌溉对冬小麦耗水特性、光合特性、产量及水分利用效率(WUE)的影响.结果表明,小麦播种至拔节期,以消耗0-40 cm土层水分为主,在此期间42 mm降水条件下,不灌冻水处理在拔节前0-40 cm土层达到重度水分亏缺,灌冻水处理只为轻度水分亏缺.前期重度水分亏缺对后期根系吸收深层水分和旗叶光合速率起到显著影响.轻度水分亏缺条件下,气孔导度下降,蒸腾速率随之降低;而光合速率可得以维持,单叶水分利用效率提高.随灌水次数增加,总耗水量加大,土壤水和降水的消耗比例显著降低.产量、WUE与耗水量均呈二次曲线关系,但变化趋势不一致,两曲线在耗水量360 mm处相交,为两者理论上最佳结合点.本试验中,冻水+拔节水处理产量最高,达到7753 kg/hm2,比不灌水处理(WO)提高了40.2%,WUE值为1.9 ks/m3,与WO处理差异不显著,为本试验的最优节水高产灌溉方案.

... 3% ^[7] ...

... mol m^-2 s^{-1 [7]} ...

2007

0.0

1.667

. 2007, 33:1884-1891

Relationship of water use efficiency with photoassimilate accumulation and transport in wheat under deficit irrigation

(亏缺灌溉下小麦水分利用效率与光合产物积累运转的相关研究) . (作物学报) , 2007 , 33 : 1884 - 1891 Hu M Y , Zhang Z B , Xu P , Dong B D , Li W Q , Li J J . Relationship of water use efficiency with photoassimilate accumulation and transport in wheat under deficit irrigation

Hu M Y

(胡梦芸) , (张正斌) , (徐萍) , (董宝娣) , (李魏强) , (李景娟) . (亏缺灌溉下小麦水分利用效率与光合产物积累运转的相关研究) . (作物学报) , 2007 , 33 : 1884 - 1891 Hu M Y

1 Center of Agricultural Resources Research, Institute of Genetic and Developmental Biology, Chinese Academy of Sciences, Shijiazhuang 050021, Hebei; 2 Graduate University of Chinese Academy of Sciences, Beijing 100049, China

Deficit irrigation in winter wheat (Triticum aestivum L.) has been practiced in the areas with limited irrigation water resources. The objectives of this study were to (i) investigate the effect of different deficit irrigations on yield and water use efficiency (WUE); and (ii) understand the physiological mechanisms of drought resistance, water saving and high yield under deficit irrigation in wheat. A field experiment was conducted using four wheat (Jinmai 47 and Xifeng 20 in dryland, Shijiazhuang 8 and 4185 in dry-wet land) with the treatments of rainfed (T0), watering once at jointing (T1) and watering twice at jointing and anthesis (T2). The amount of irrigation is 60 mm each time. The physiological parameters measured included net photosynthetic rate (P_n), stomatal conductance (G_s), leaf area index, dry matter accumulation and partitioning, root distribution, yield components and water consumption, as well as the corelation between WUE and yield components. Non-irrigation before jointing and deficit irrigation from jointing to anthesis could significantly improve dry matter production and deep root development, and significantly save irrigation water. Both grain yield and WUE were conically related with water consumption. Non-irrigation significantly decreased dry matter accumulation and number of spikes. Low available soil water from booting to late grain filling resulted in the lowest biomass, grain yield, harvest index and WUE in T0 treatment. The increased yield in T1 and T2 was mainly contributed by the increased number of spikes, and seeds per spike. The increased WUE was mainly contributed by increased HI. Water stress during grain filling reduced Pn and G_s, and accelerated photosynthetic leaf senescence. However, the water stress during grain filling induced remobilization of pre-anthesis carbon reserves to grains, significantly contributing the increased grain yield and HI. It is concluded that deficit irrigation between jointing and anthesis can change the root spread and develop deep root system, and significantly increase soil water use efficiency. Increasing both photosynthesis and photosynthetic duration, as well as increasing the remobilization of pre-anthesis carbon reserves and HI must increase wheat yield and WUE.

在大田栽培条件下，以小麦旱地品种晋麦47和西峰20、水旱兼用型品种石家庄8号和水地品种4185为材料，分别进行0水（T0）、一水（T1）和二水灌溉（T2）处理（每次灌水量60 mm），研究了光合速率、叶面积指数、干物质积累与分配、根系分布、耗水量、产量因子与水分利用效率（WUE）的关系。结果表明，在拔节前不灌溉，拔节到开花期亏缺灌溉，促进干物质积累和深根发育。随着灌溉水的增加，耗水量显著增加，产量和WUE与耗水量呈二次曲线关系。T0处理显著减少了干物质积累和成穗数，产量、经济系数（HI）和WUE最低。T1和T2产量的提高主要是增加了穗数和穗粒数。灌浆期水分亏缺降低了光合速率（P_n）和气孔导度（G_s），加速了功能叶片的衰老，但诱导了花前储存碳库的再转运，显著提高了HI和产量。因此，在拔节和开花期亏缺灌溉促进根系生长，提高了土壤水分的利用效率。而产量和产量WUE的提高主要是由于增加了灌浆期叶片的P_n和光合功能持续期，促进花前储存碳库的再转运，显著提高了HI。

... 6% ^[8] ...

... 各指标计算公式^[8]如下 ...

... 3% ^[8] ...

... 16 g, 且花前干物质积累及向籽粒再转运量较大, 对籽粒灌浆贡献率为47% ^[8] ...

2011

0.0

0.795

. 2011, 19:805-811

Effect of limited irrigation on diurnal variation in flag-leaf photosynthesis and yield of dryland wheat

(限量灌溉对旱地小麦旗叶光合特性日变化和产量的影响) . (中国农业生态学报) , 2011 , 19 : 805 - 811 Tan N T , Lin Q , Jiang W , Liu Y G , Li L Y . Effect of limited irrigation on diurnal variation in flag-leaf photosynthesis and yield of dryland wheat

Tan N T

(谭念童) , (林琪) , (姜雯) , (刘义国) , (李玲燕) . (限量灌溉对旱地小麦旗叶光合特性日变化和产量的影响) . (中国农业生态学报) , 2011 , 19 : 805 - 811 Tan N T

A field experiment was conducted to study the effects of five irrigation treatments on diurnal variations in flag-leaf photosynthesis and yield of “Qingmai 6” dryland wheat variety. The results showed that diurnal variations in net photosynthetic rate (Pn) in each treatment followed a double-peak curve. However, the degree of midday depression in photosynthesis increased with increasing amount of irrigation. Increasing irrigation amount enhanced flag-leaf Pn and net stomatal conductance (Gs) under ≤3 irrigation times. The maximum Pn and Gs appeared in W3 treatment (i.e., 60 mm irrigation each at jointing, booting and grain-filling stages). With increasing supplementary irrigation, intracellular CO₂ concentration and stomatal limitation (Ls) decreased. Flag-leaf Pn under W4 treatment (i.e., 60 mm irrigation each at sprouting, jointing, booting and grain-filling stages) was significantly lower than that under W3 treatment. This suggested that superfluous irrigation had negative effects on flag-leaf photosynthesis in dryland winter wheat at grain-filling stage. In all the treatments, yield under W2 treatment (i.e., 60 mm irrigation each at jointing and booting stages) was the highest. The difference in yield between W2 and W1 (i.e., 60 mm irrigation at jointing stage) treatments was insignificant. Under ≤2 times of irrigation, yield increased with increasing irrigation amount. Yields under W3 and W4 treatments were lower than that under the dryland control treatment. This suggested that superfluous irrigation was unsuitable for high yield dryland winter wheat cultivation. In all the treatments, water production efficiency (WPE) declined with increasing irrigation amount. The experiment suggested that irrigation at jointing, booting and grain-filling stages was crucial for high flag-leaf Pn in dryland winter wheat. However, the effect of irrigation at sprouting stage on flag-leaf photosynthesis of dryland winter wheat was insignificant. Jointing was the most pivotal stage for high yield of wheat. Jointing and booting were the most essential stages for irrigating of winter wheat. Dryland wheat irrigation at these stages was very important for high yield and WPE. In terms of yield and WPE, the optimal supplementary irrigation mode for high yield in dryland winter wheat was 60 mm at jointing stage.

在大田条件下, 以“青麦6 号”小麦为试验材料, 研究了5 个灌水处理下旱地冬小麦灌浆期旗叶光合特性日变化和产量的变化。结果表明, 5 个灌水处理净光合速率日变化曲线均为双峰曲线, 但随着灌水次数的增加, “午休”明显减弱。在灌溉低于3 水的情况下, 旗叶的净光合速率、气孔导度都随灌溉量的增加呈上升趋势,均以灌3 水(W3, 拔节水60 mm+孕穗水60 mm+灌浆水60 mm)处理最高, 而胞间CO₂ 浓度和气孔限制值随着灌溉量的增加呈下降趋势; 而灌溉4 水(W4, 起身水60 mm+拔节水60 mm+孕穗水60 mm+灌浆水60 mm)处理的净光合速率明显小于W3 处理, 表明过量灌溉对旱地小麦灌浆期旗叶的光合作用有消极作用。小麦产量以灌2 水(W2, 拔节水60 mm+孕穗水60 mm)处理为最高, 且在灌溉2 水以下随着灌溉次数增加而增加, 但灌1 水(W1, 拔节水60 mm)和W2 处理间产量差异不显著, 同时灌溉3 水(W3)和4 水(W4)会使产量下降, 反而低于旱地处理, 因此过多灌水不利于旱地小麦的高产。试验结果还表明, 拔节期、孕穗期和灌浆期灌溉对旱地小麦旗叶灌浆中期达到较高光合作用至关重要, 而起身水对旱地小麦光合作用无显著影响。拔节期是旱地小麦达到高产最重要的灌溉时期, 拔节~孕穗期为冬小麦需水关键期, 拔节~孕穗水对冬小麦产量和水分利用效率有重要影响。水分生产效率也随着灌水量的增加呈下降趋势。综合产量和水分利用效率因素, 以灌溉拔节水60 mm(W1)处理为达到旱地冬小麦高产的最佳灌溉模式。

... mol m^-2 s^{-1 [9]} ...

2009

0.0

1.667

. 2009, 35:2045-2054 DOI:10.3724/SP.J.1006.2009.02045

Utilization of water and nitrogen and yield formation under three limited irrigation schedules in winter wheat

(冬小麦节水栽培三种灌溉模式的水氮利用与产量形成) . (作物学报) , 2009 , 35 : 2045 - 2054 Zhang S Q , Fang B T , Zhang Y H , Zhou S L , Wang Z M . Utilization of water and nitrogen and yield formation under three limited irrigation schedules in winter wheat

Zhang S Q

(张胜全) , (方保停) , (张英华) , (周顺利) , (王志敏) . (冬小麦节水栽培三种灌溉模式的水氮利用与产量形成) . (作物学报) , 2009 , 35 : 2045 - 2054 Zhang S Q

1College of Agronomy and Biotechnology,China Agricultural University,Beijing 100193,China;2Wheat Research Center,Henan Academy of Agricultural Sciences,Zhengzhou 450002,China

Water shortage is a great challenge in winter wheat (Triticum aestivum L.) production worldwide. Studies on wheat physiology and ecology have been carried out aiming at yielding more grains with less consumptions of water and nitrogen fertilizer. Cultivation techniques based on limited irrigation and nitrogen application are highly emphasized in North China Plain, where water stress is the most major problem in regular years. In Wuqiao area, Hebei province, China, a series cultivation systems have been established with distinct effects of high yield and small water consumption. The principle measure in the technique system was controlling the irrigation frequency and water amount. The purpose of this study was to disclose the mechanism of high-yielding and high resource use efficiency in the cultivation system established in Wuqiao. In the consecutive growing seasons from 2004 to 2008, field experiments were conducted at the Wuqiao Experimental Station of China Agriculture University under three limited irrigation schedules, i.e., no irrigation in spring (I₀), 750 m³ha^-1 of water at jointing (I₁) and each 750 m³ha^-1 of water at jointing and anthesis (I₂). Fertilizers were only applied before sowing, including urea 225 kg ha^-1, (NH₄)₂HPO₄ 300 kg ha^-1 (thus, the total nitrogen was 157.5 kg ha^-1),K₂SO₄225 kg ha^-1, and ZnSO₄22.5 kg ha^-1. The characteristics of water and nitrogen use and the formation of grain yield were investigated at jointing, booting, anthesis, and maturity stages. The results showed that the average yield was 6134 kg ha^-1 for I₀, 7 515 kg ha^-1 for I₁, and 8 134 for I₂ treatment, and the average total evapotranspiration was in the order of 3 334, 3 829, and
4 270 m³ha^-1. The ratio of soil water consumption to total evapotranspiration was 67%, 49%, and 38% in I₀, I₁, and I₂ treatments, respectively. The water use efficiency ranged from 1.9 to 2.0 kg m^-3 with no significant differences among treatments. This indicated that approximately 52.6 m³ of water was required for yielding 100 kg of grains. Under the nitrogen application level of 157.5 kg ha^-1, the average total nitrogen accumulation was 163, 198, and 212 kg ha^-1 in I₀, I₁, and I₂ treatments, respectively. Consistent results with slight variations were observed in the total nitrogen accumulation across years. The average nitrogen use efficacy of the three irrigation treatments was 38 kg kg^-1and there were no significant differences among treatments. In terms of the characteristics of plant population, the leaf area index (LAI) at booting and anthesis, as well as the LAI for the top three leaves were not significantly different between I₁ and I₂ treatments, but they were significantly higher than those in I₀ treatment. The LAI for the top three leaves and non-leaf organs green area index (NAI) above flag leaf ranged from 2.6 to 3.8 and from 2.7 to 3.6, respectively. The ratios of non-leaf organs green area above flag leaf to the uppermost three leaf area and total leaf area were more than 1.0 and 0.7, respectively. Three limited irrigation schedulesshowed steady grain yield and high resource use efficiency for consecutive four years.

为探讨华北平原冬小麦节水栽培适宜灌溉模式及其产量形成与资源利用规律，2004—2008连续4年在同一地块比较春不灌水(I₀)、春灌1水(I₁)和春灌2水(I₂) 3种灌溉模式对冬小麦水氮利用特性与产量形成的影响。在均实行晚播和一次性底施氮量157.5 kg hm^-2条件下，各模式年际间籽粒产量稳定，I₀、I₁和I₂模式的4年平均产量分别为6 134、7 515和8 134 kg hm^-2，平均总耗水量分别为3 334、3 829和4 270 m³ hm^-2,平均总吸氮量分别为162.5、197.9和212.2 kg hm^-2。在耗水构成中，土壤贮水消耗占总耗水量的比例分别为66.7%、49.2%和37.8%。3种灌溉模式的水分利用效率达1.9~2.0 kg m^-3，灌溉模式间无显著差异；氮素利用效率在3个灌溉模式间也无显著差异，平均为38.0 kg kg^-1 N。3种灌溉模式主茎穗占成穗数比例平均为81%，开花期上三叶叶面积指数2.6~3.8，旗叶节以上非叶绿色器官面积指数2.7~3.6。综合以上结果，冬小麦3种灌溉模式及其配套技术形成的群体结构可在不同产量水平上实现水、氮资源高效利用。

... 4 g ^[10] ...

... 2 ^[10] ...

2012

0.0

... 7 kg hm^{-2 [11]} ...

2004

0.0

... _j) ^[12], 式中, M为灌水量(mm), H为土壤计划湿润层的深度(cm), #cod#x003b3 ...

2004

0.0

1.299

. 2004, 20(4):58-63

Research on the water use efficiency and optimal irrigation schedule of the winter wheat

(冬小麦水分利用效率与最优灌溉制度的研究) . (农业工程学报) , 2004 , 20 ( 4 ): 58 - 63 Liu Z J , Li B P , Li Y H , Cui Y L . Research on the water use efficiency and optimal irrigation schedule of the winter wheat

Liu Z J

(刘增进) , (李宝萍) , (李远华) , (崔远来) . (冬小麦水分利用效率与最优灌溉制度的研究) . (农业工程学报) , 2004 , 20 ( 4 ): 58 - 63 Liu Z J

针对不同水分处理对冬小麦产量和耗水特性的影响,在大田生产条件下对冬小麦进行了非充分灌溉试验,以冬小麦大田试验数据为基础,采用最小二乘法拟合了冬小麦水分生产函数;通过耗水资料的分析整理,揭示了冬小麦的水分效应及耗水规律;依据动态规划原理,建立了冬小麦最优灌溉制度动态模型,提出了两种水文年型(50%和75%)的冬小麦节水灌溉模式,对北方冬麦区田间管理具有一定的指导意义.

... S+M+P+K ^[13,14] ...

2013

2.8

0.0

... S+M+P+K ^[13,14] ...

2007

0.0

... 本试验地区地下水埋深在4 m以下^[15], 因此无地下水补给 ...

2011

2.289

0.0

. 2011, 29:213-232 DOI:10.1007/s00271-010-0229-1

Differences in water-use efficiency among perennial forages used by the dairy industry under optimum and deficit irrigation

1.Industry and Investment NSW, Elizabeth Macarthur Agricultural Institute, Woodbridge Rd, Menangle, New South Wales 2570, Australia<br/>2.Faculty of Veterinary Science, The University of Sydney, MC Franklin Lab, Camden, New South Wales 2570, Australia<br/>3.Faculty of Agriculture, The University of Sydney, Food and Natural Resources, Sydney, New South Wales 2006, Australia<br/>

The cost and scarcity of water is placing increasing pressure on Australian dairy farmers to utilise water for forage production as efficiently as possible. This study aimed to identify perennial forage species with greater water-use efficiency (WUE) than the current dominant species, perennial ryegrass (Lolium perenne L.). Fifteen perennial forage species were investigated under optimum irrigation and two deficit irrigation treatments, over three years at Camden, NSW, on a brown Dermsol in a warm temperate climate. Under optimal irrigation, there was a nearly twofold difference in mean WUE_t (total yield/evapotranspiration) between forages, with kikuyu (Pennisetum clandestinum Hochst. ex. chiov.) having the highest (27.3 kg ha⁻¹ mm⁻¹) and birdsfoot trefoil (Lotus corniculatus L.) the lowest (14.8 kg ha⁻¹ mm⁻¹). Kikuyu was also the most water use efficient forage under the extreme deficit irrigation treatment, although its mean WUE_t declined by 15% to 23.2 kg ha⁻¹ mm⁻¹, while white clover (Trifolium repens L.) in the same treatment had the largest decline of 44% and the lowest WUE_t of only 8.8 kg ha⁻¹ mm⁻¹. In order to maximise WUE for any forage, it is necessary to maximise yield, as there is a strong positive relationship between yield and WUE_t.

... 水分利用效率 = 籽粒产量 / 作物全生育期耗水量^[16,17] ...

2012

0.0

... 水分利用效率 = 籽粒产量 / 作物全生育期耗水量^[16,17] ...

2009

1.113

0.0

... 依据Li等的方法^[18], 叶面积 = 最大叶长#cod#x000d7 ...

2007

0.0

1.667

. 2007, 33:1267-1271

Diurnal changes of photosynthetic and physiological parameters in different male sterile lines of wheat

(不同小麦雄性不育类型光合、生理参数日变化的研究) . (作物学报) , 2007 , 33 : 1267 - 1271 Jiao J , Gao Q R , Zhang A M , Hao Y Y , Wang D W , Wang L , Qiu X M . Diurnal changes of photosynthetic and physiological parameters in different male sterile lines of wheat

Jiao J

(焦健) , (高庆荣) , (张爱民) , (郝媛媛) , (王大伟) , (王霖) , (邱新民) . (不同小麦雄性不育类型光合、生理参数日变化的研究) . (作物学报) , 2007 , 33 : 1267 - 1271 Jiao J

1 Agronomy College, Shandong Agricultural University/Key Laboratory of Corp Biology of Shandong Province, Tai’an 271018, Shandong; 2 Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101; 3 Jining Academy of Agricultural Science, Jining 272031, Shandong, China

In order to study the cytoplasmic effects of male sterile wheat, male sterile lines of CHA (chemical hybridizing agent) and cytoplasmic K (Aegilops kotschyi), T (Triticum timopheevi), V (Aegilops ventricosa) types of male sterile wheat at different developmental stages were compared with their maintainer lines in the diurnal changes of photosynthetic and physiological parameters, comprising net photosynthetic rate (P_n), stomatal conductance (G_s), transpiration rate (T_r), water use efficiency (WUE), photon flux density (PFD), intercellular CO₂concentration (C_i), atmospheric CO₂ concentration (C_a), leaf temperature (T_i), atmospheric temperature (T_a), and stomatal limiting value (L_s). And these parameters may reflect the photosynthetic efficiency of the leaves and environmental adapting capability of the plant. The experiment was conducted to analyze the diurnal changes and the physiological mechanism of photosynthetic midday-depression, the response to male sterile cytoplasms of female parent genotypes and the differences of photosynthetic and physiological parameters of 4 male sterile lines and their maintainer lines, which can reflect differences of heredity of 4 male sterile cytoplasms under the same nuclear background and also exhibit differences of the effect of nuclear-cytoplasmic interaction. The result indicated that, the curve of diurnal changes of P_n presented two peak values, with one peak at 10:00 higher than the other at 14:00. And the phenomenon of photosynthetic midday-depression of leaf happened at noon. The diurnal changes of Pn was positively correlated with G_s, C_i and negatively correlated with L_s, and the correlation coefficients were 0.8187*, 0.8136*, and －0.8496* respectively. Accompanied by the decline of G_s, C_i and the raise of L_s, P_n declined distinctly in midday, which means the limitation of Pn by stomatal factor. Compared with their maintainer lines, CHA and K, T, V-male sterile lines had lower P_n values, decreasingly 0.88, 2.76, 3.30, and 2.04 μmol CO₂ m^-2 s^-1 respectively, indicating that the CHA and K, T, V-cytoplasm has negative effects on Pn with different extent, especially T and K-types. K, T, V cytoplasmic male sterile lines had lower WUE values than their maintainer lines, decreasingly 0.94–1.54 μmol CO₂ mmol^-1 H₂O, and indicating that K, T, V-cytoplasm has significantly negative effects on WUE. Compared with the maintainer lines, CHA-sterile lines has no significant difference in WUE values, indicating chemical hybriding agent (BAU-9403) has no significantly negative effects on the water use efficiency of wheat flag leaf. The influences of damage caused by BAU-9403 were less than the negative effects of other male sterile cytoplasms. Moreover, the Pn value varied between genotypes of different female parents, Pn of Ji 5418 and its CMS lines showed significant or highly significant differences with that of Tai 911289, indicating that genotypes of female parents have different effects on different cytoplasms in Pn. This result also demonstrates the enormous potential for improving the photosynthetic capability in Ji 5418. Therefore, it is better to reduce the negative effects of male sterile cytoplasms on photosynthetic function of wheat by selecting preponderant recurrent parent to make backcross in a breeding project.

利用2套同核异质的K、T、V、CHA型不育系及其保持系，对其不同生育时期净光合速率（P_n）、气孔导度（G_s）、蒸腾速率（Tr）、水分利用率（WUE）、光量子通量密度（PFD）和细胞间隙CO₂浓度（C_i）、气孔限制值（L_s）等光合、生理参数的日变化及其差异进行比较研究，探讨不同小麦雄性不育类型的胞质效应。结果表明，供试材料净光合速率的日变化均呈双峰曲线，有明显的光合“午休”现象，P_n与G_s、C_i的相关系数分别为0.8187、0.8136，呈显著正相关（r_0.05=0.8110），与L_s的相关系数为－0.8496，呈显著负相关（r_0.05=0.8110），P_n的午间下降伴随着G_s、Ci的下降和L_s的升高，表明其主要受气孔因素限制；与保持系的Pn值相比，CHA型和K、T、V型不育系分别降低0.88、2.76、3.30和2.04 μmol CO₂ m^-2 s^-1，产生不同程度的负效应，尤以T型和K型最明显。在水分利用率上，K、T、V型不育系较保持系降低0.94~1.54 μmol CO₂ mmol^-1 H₂O，负效应显著；CHA型不育系较保持系低0.36 μmol CO₂mmol^-1 H₂O，差异不显著，说明CHA对旗叶水分利用率无明显的不良影响。Pn值因母本基因型的不同而异，冀5418及相应不育系的Pn值较太911289的高，差异达显著或极显著水平。因此，选择优势效应大的保持系回交，有利于降低不育胞质对小麦光合功能的负效应。

... 气孔限制值L_s= 1-C_i/C_a^[19] ...

2004

2.853

0.0

... 4 ^[20] ...

2006

0.0

1.667

. 2006, 32:70-75

Stomatal characteristics of different green organs in wheat under different irrigation regimes

(不同供水条件下小麦不同绿色器官的气孔特性研究) . (作物学报) , 2006 , 32 : 70 - 75 Zhang Y P , Wang Z M , Wu Y C , Zhang X . Stomatal characteristics of different green organs in wheat under different irrigation regimes

Zhang Y P

(张永平) , (王志敏) , (吴永成) , (张霞) . (不同供水条件下小麦不同绿色器官的气孔特性研究) . (作物学报) , 2006 , 32 : 70 - 75 Zhang Y P

College of Agronomy and Bio-technology, China Agricultural University, Beijing 100094

Stomata are the channel of CO₂ and water vapour exchange between the plant and atmosphere, and contributing to photosynthesis, transpiration and temperature fitness of plant. There have been many reports about the stomatal structure and characteristics focused on the leaf blades, the major photosynthetic organs, in wheat but only a few reports about stomatal distribution on the ear（especially on the awn）, whereas, no any report on stomatal structure characteristics and their relations to photosynthetic rate and transpiration rate of various non-leaf green organs in wheat under different water condition. In current study, a field experiment was conducted with two cultivars (Lumai 21 and SJZ 8) and three irrigation treatments[No irrigation, Two irrigations (1 500 m³∙hm^-2) and Four irrigations (3 000 m³∙hm^-2) in the growing period of wheat], in order to asses differences in stomatal structure characteristics and their response to irrigation between flag leaf blade and non-leaf green organs (flag leaf sheath, peduncle, glume, lemma, palea and awn). Stomatal structure and distribution on leaf blade and non-leaf green organs in winter wheat were observed, and the stomatal conductance, transpiration rate and photosynthetic rate were also measured in response to different supplemental irrigation treatments. It was showed that under different irrigation treatments, stomata could be found on all surfaces of flag leaf blade , sheath and peduncle, and also on the surface of awn and abaxial surface of glume in ear. However, for lemma, stomata appeared only on the adaxial (not on the abaxial )side under both two and four irrigation treatments, and only on the abaxial side under water stress(no irrigation). In size, stomat apparatus on the ear（glume, lemma, palea and awn）was smaller than that on other organs. With the decrease of water supply, the width of stomatal aperture was also decreased, while both stomat frequency and stomatal aperture length of various organs were increased. Compared with flag leaf blade, non-leaf green organs (flag leaf sheath, peduncle and ear) was less sensitive to supplemental irrigation, and showed higher stability in stomatal conductance, transpiration rate and photosynthetic rate during the later filling period. There was a positive correlation between stomatal conductance and transpiration rate in various organs, while the coefficient of correlation in non-leaf green organs was lower than that in leaf blade. The results suggested that with the decrease of water supply, transpiration rates were declined for both the leaf blades and non-leaf green organs, then, photosynthetic rates might be remarkably decreased in leaf blades, however, relatively stable in non-leaf green organs, which may lead to higher water use efficiency (WUE) in non-leaf green organs than in leaf blades.

为了考察小麦叶片与非叶器官气孔结构特性的差异及其对供水条件的反应，本研究设置不同灌水处理，利用电镜观察小麦灌浆期不同绿色器官的气孔分布和结构特征，并分析其与气孔特性指标间的关系。结果表明，在不同灌水处理下看到各非叶器官（穗、旗叶鞘和穗下节间）均分布着气孔，但其数目少于旗叶叶片。护颖仅在远轴面存在气孔；外稃在多水条件下（4水处理）近轴面出现较多气孔，而远轴面看不到气孔，但在水分胁迫（无水处理）条件下，气孔却出现在远轴面而不在近轴面；在不同水分处理下均观察到芒上明显的气孔分布。从气孔大小看，穗各部分（护颖、外稃、内稃和芒）略小于其他器官。随着灌水次数的减少，各器官气孔密度呈增大趋势，气孔器及气孔孔径表现出长度增加、宽度减小的特征。限水灌溉下非叶器官（穗、旗叶鞘和穗下节间）在籽粒灌浆期气孔导度、蒸腾速率和光合速率的稳定性高于叶片。相关分析表明，不同器官的气孔导度与蒸腾速率均呈显著正相关，非叶器官气孔导度与光合速率的相关程度明显低于叶片。说明在干旱少水条件下，叶与非叶器官蒸腾作用均会减弱，叶片光合速率亦相应降低，而非叶器官光合速率可能保持相对稳定，可相对提高其水分利用效率。

... 不同供水条件下, 随灌水量减少各器官气孔密度增大, 其中旗叶增幅较大, 约9%, 与气孔导度呈负相关^[21], 导致气孔限制值升 ...

2012

0.0

1.667

. 2012, 38:1062-1070 DOI:10.3724/SP.J.1006.2012.01062

Effects of partial root zone irrigation on leaf photosynthetic curves and chlorophyll fluorescence parameters in naked oat

(局部根区灌溉对裸燕麦光合特征曲线及叶绿素荧光特性的影响) . (作物学报) , 2012 , 38 : 1062 - 1070 Lin Y C , Zeng Z H , Ren C Z , Li Z J , Guo L C , Yang X C , Wang C L , Qian X , Hu Y G . Effects of partial root zone irrigation on leaf photosynthetic curves and chlorophyll fluorescence parameters in naked oat

Lin Y C

(林叶春) , (曾昭海) , (任长忠) , (李志坚) , (郭来春) , (杨学超) , (王春龙) , (钱欣) , (胡跃高) . (局部根区灌溉对裸燕麦光合特征曲线及叶绿素荧光特性的影响) . (作物学报) , 2012 , 38 : 1062 - 1070 Lin Y C

<div style="line-height: 150%">1College of Agronomy and Biotechnology, China Agricultural University, Beijing 100193, China; 2Baicheng Academy of Agricultural Sciences, Baicheng 137000, China; 3Institute of Grassland Science, Northeast Normal University, Changchun 130024, China</div>

The objective of this study was to provide experimental evidence for photosynthetic mechanism under drought stress and practical technique of water-saving irrigation in naked oat (Avena nuda L.). In a pot experiment with instrument to control root zone irrigation and water amount, the effects on relative chlorophyll content (SPAD), photosynthetic curves, and chlorophyll fluorescence parameters were compared among irrigation models of alternative partial root zone irrigation (APRI), fixed partial root zone irrigation (FPRI), and conventional total root zone irrigation (CTRI). Compared to CTRI, partial root zone irrigation, including APRI and FPRI treatments, decreased leaf SPAD, net photosynthetic rate (P_n), and carboxylation efficiency (CE); however, APRI had no obvious effects on reducing initial quantum efficiency (α), maximum photochemical efficiency (F_v/F_m), actual photochemical efficiency of PSII (Ф_PSII), apparent electron transport rate (ETR), and photochemical quenching coefficient (q_P). The leaf SPAD of APRI was significantly enhanced compared to that of FPRI (P<0.05), and the maximum net photosynthetic rate (P_max), α, light saturation point (LSP), light use efficiency (LUE), C_i/C_a, CE, CO₂ saturation point (C_i,sat), F_o, F_m, Ф_PSII, ETR, q_P, and NPQ were all higher in APRI than in FPRI. The reduction of P_n in partial root zone irrigation treatments resulted from stomatal factors, and structural damage of photosynthetic system II (PSII) was found in FPRI treatment. Clearly, partial root zone irrigation can improve the tolerance ability of naked oat against drought stress; particularly, APRI has the effect to maintain higher P_n than FPRI.

为探讨局部根区不同灌溉方式下裸燕麦(Avena nuda L.)光合能力下降的生理机制，采用盆栽及渗水盘供水方法，比较了交替根区灌溉(APRI)、固定根区灌溉(FPRI)和常规灌溉(CTRI)下，裸燕麦旗叶相对叶绿素含量(SPAD值)、光合特征曲线及叶绿素荧光动力学特性的差异。与CTRI处理相比，局部根区灌溉(包括APRI和FPRI处理)降低了叶片SPAD值、净光合速率(P_n)和初始羧化速率(CE)，但APRI处理未明显降低初始量子效率(α)、PSII最大量子效率(F_v/F_m)、PSII实际光化学效率(Ф_PSII)、电子传递效率(ETR)和光化学效率 (q_P)。在2种局部根区灌溉模式中，APRI较FPRI显著提高了叶片SPAD值(P<0.05)，而且APRI的叶片最大净光合速率(P_max)、α、光饱和点(LSP)、光能利用率(LUE)、C_i/C_a、CE、CO₂饱和点(C_i,sat)、初始荧光(F_o)、最大荧光(F_m)、Ф_PSII、ETR、q_P和非光化学效率(NPQ)均高于FPRI。APRI和FPRI的光合速率降低与气孔因素有关，FPRI光合速率降低还与PSII结构损伤有关；局部根区灌溉提高了裸燕麦干旱胁迫逆境下的耐受能力，APRI有利于保持更高的光合速率。

... mol m^-2 s^{-1 [22]} ...

2013

0.0

... Zhao等^[23]研究指出, 拔节期和抽穗期分别灌溉60 mm, 小麦抽穗期叶面积指数显著提高, 达到8 ...

2010

0.0

. 2010, 30:1548-1555

Effects of irrigation frequency and stages on grain yield and quality characteristics of winter wheat

(灌溉频次和时期对冬小麦籽粒产量及品质特性的影响) . (生态学报) , 2010 , 30 : 1548 - 1555 Han H F , Li Q Q , Dong B D , Liu M Y . Effects of irrigation frequency and stages on grain yield and quality characteristics of winter wheat

Han H F

(韩慧芳) , (李全起) , (董宝娣) , (刘孟雨) . (灌溉频次和时期对冬小麦籽粒产量及品质特性的影响) . (生态学报) , 2010 , 30 : 1548 - 1555 Han H F

为探讨我国北方地区冬小麦的节水灌溉模式,2006-2008年,在中国科学院栾城农业生态系统试验站,以冬小麦品种科农9204为试验材料,在总灌溉量为120mm的条件下,研究了灌溉次数和灌溉时期对籽粒产量、水分利用效率(WUE)、籽粒蛋白质含量以及相关主要品质特性的影响.结果表明,拔节期、抽穗期和灌浆期一次灌溉分别有利于产量、干物质积累量和千粒重的形成或提高;两次灌溉处理中,以拔节和抽穗期各灌60mm处理籽粒产量最高,籽粒蛋白质产量有随灌溉时期后移而降低的趋势;冬小麦生育期间随灌溉次数增多和灌溉时期后移,湿面筋含量、面团形成时间、面团稳定时间等均显著降低.综合考虑冬小麦的籽粒产量、WUE、营养品质和加工品质,在总灌溉量为120mm的条件下,以拔节和抽穗期各灌溉60mm为宜.

Abstract：

In North China,in order to obtain high and stable grain yield of winter wheat,irrigation should be applied during its growing season.However,water resource is very shortage and irrigation water use efficiency is very low.Therefore,irrigation frequency should be adopted in this region.In 2006-2008,an experiment was conducted at Luancheng Experimental Station of CAS to study irrigation frequency on grain yield,water use efficiency (WUE),protein yield,wet gluten content,development time,and stability time of winter wheat under irrigation frequency by variety Kenong "9204".The results showed that irrigated at jointing,heading,and milking stages were in favor of increased grain yield,dry matter accumulation,or 1000 kernel weight,respectively.The highest grain yield was obtained in case of the treatment was irrigated 60 mm at jointing and heading stages,respectively.Irrigated two times during the growing stages of winter wheat could increase protein yield:however,irrigation timing during the later part of the growing season of winter wheat could decrease the protein yield.The results indicated that irrigation timing in the later part of the growing season and the increase in the times of irrigation,wet gluten content,development time,and stability time were all significantly decreased.On the basis of the experimental results,it is suggested that irrigation should be applied at jointing and heading stages of winter wheat to obtain reasonable grain yield,WUE,and quality under irrigation frequency in North China.

... 1% ^[24] ...

2001

1.67

0.0

. 2001, 35:81-91 DOI:10.1023/A:1013827600528

The effect of water stress on the activities of key regulatory enzymes of the sucrose to starch pathway in wheat

1.University of London<br/>

Developmental changes in the starch and sucrose content of grains andthe activities of enzymes of starch synthesis in wheat were studied under waterstress conditions. Water stress caused a marked reduction in the sucrose andstarch content of the grains. Sucrose synthase (SS) and UDP-glucosepyrophosphorylase (UDP-Gppase), showed higher catalytic activity and moreresistance to water stress compared with amyloplastic enzymes. ADP-glucosepyrophosphorylase (ADP-Gppase) activity was reduced to a low level under bothin situ and osmotic stress conditions in which grainsfailed to accumulate dry matter in vivo. Granule-boundstarch synthase (GBSS) also responded rapidly to in situwater stress treatments as did ADP-Gppase. Reduction in GBSS activity at thetime of growth cessation in situ was less than that ofADP-Gppase and the enzyme did not respond to severe osmotic stress. Solublestarch synthase (SSS) was the enzyme most sensitive to water stress in that itresponded earlier, and to a greater extent, than the other enzymes. However,under severe dehydration conditions, leading to cessation of growth, thedeclinein SSS activity was less than that for ADP-Gppase. SSS showed the lowestin vitro activity followed by GBSS. These results suggestthat SSS is the site of response to water stress by which the rate of graingrowth can be affected, whereas growth cessation is due mainly to theinactivation of ADP-Gppase.

... 与水分充足处理相比, 小麦开花后水分亏缺, 导致开花后16 d籽粒中同化物降低22%, 开花后32 d降低48%^[25] ...

2002

0.0

... 灌浆后期, 70%以上同化物运送至籽粒中^[26] ...

2012

0.0

... 亦有研究表明, 花前累积干物质对小麦产量有一定贡献, 但产量高低主要取决于花后干物质的积累和转移, 花后转移干物质贡献率可达65%以上^[27] ...

2011

1.265

0.0

... 4 kg m^{-3 [28]} ...

2008

0.0

... 42 kg m^{-3 [29]} ...