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作物学报 ›› 2023, Vol. 49 ›› Issue (6): 1653-1667.doi: 10.3724/SP.J.1006.2022.21045

• 耕作栽培·生理生化 • 上一篇    下一篇

灌溉对小麦分蘖发生和不同茎蘖光合同化物生产分配及成穗的影响

冯连杰(), 于振文, 张永丽*(), 石玉   

  1. 山东农业大学农学院/作物生物学国家重点实验室/农业农村部作物生理生态与耕作重点实验室, 山东泰安 271018
  • 收稿日期:2022-06-28 接受日期:2022-10-10 出版日期:2023-06-12 网络出版日期:2022-12-26
  • 通讯作者: *张永丽, E-mail: zhangyl@sdau.edu.cn
  • 作者简介:E-mail: 18853866772@163.com
  • 基金资助:
    泰山学者工程专项;财政部和农业农村部国家现代农业产业技术体系建设专项(CARS-3-1-19);国家自然科学基金项目(31771717)

Effects of irrigation on tiller occurrence, photo-assimilates production and distribution in different stem and tillers and spike formation in wheat

FENG Lian-Jie(), YU Zhen-Wen, ZHANG Yong-Li*(), SHI Yu   

  1. College of Agronomy, Shandong Agricultural University/State Key Laboratory of Crop Biology/Key Laboratory of Crop Ecophysiology and Farming System, Ministry of Agriculture and Rural Affairs, Tai’an 271018, Shandong, China
  • Received:2022-06-28 Accepted:2022-10-10 Published:2023-06-12 Published online:2022-12-26
  • Contact: *E-mail: zhangyl@sdau.edu.cn
  • Supported by:
    Special funds for Taishan Scholars Project;China Agriculture Research System of MOF and MARA(CARS-3-1-19);National Natural Science Foundation of China(31771717)

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摘要：

依据土壤墒情适量灌溉可增加小麦穗数, 实现节水高产, 但该灌溉条件下分蘖发生和成穗的生理机制尚不明确。本试验于2019—2020和2020—2021两个小麦生长季, 以中穗型品种济麦22和大穗型品种山农23为试验材料, 设置全生育期不灌溉、节水灌溉和充分灌溉3个处理, 研究了不同穗型小麦分蘖发生和成穗规律。结果表明, 2个品种节水灌溉处理的分蘖节面积和反式玉米素含量、越冬期和返青期主茎最上部展开叶光合参数均高于不灌溉处理, 促进了II、III、IP、IV和其余蘖发生, 济麦22和山农23单株总茎蘖数比不灌溉处理越冬期两年度平均增加1.01个和0.75个, 拔节期分别增加0.71个和0.56个。节水灌溉处理下2个品种拔节期各茎蘖最上部展开叶光合参数、干物质重和13C同化物分配量均高于不灌溉处理, 促进了分蘖成穗, 济麦22和山农23的单株成穗数比不灌溉处理两年度平均增加0.36和0.41个, 籽粒产量增加35.00%和44.27%, 水分利用效率增加9.23%和8.55%。增加灌水至充分灌溉处理, 2个品种越冬期和拔节期单株总茎蘖数较节水灌溉处理增加, 但单株成穗数、公顷穗数和籽粒产量无显著变化, 水分利用效率降低。品种间比较, 各灌溉处理下济麦22的单株总茎蘖数和单株成穗数均高于山农23, 亦归因于济麦22较高的分蘖节面积和反式玉米素含量、各茎蘖较高的光合能力。相关性分析表明, 2个品种单株总茎蘖数、单株成穗数、公顷穗数和籽粒产量与分蘖节总面积和反式玉米素含量、主茎最上部展开叶净光合速率呈正相关, 与生长素和脱落酸含量呈负相关。说明节水灌溉通过适当增加分蘖节面积和激素含量、提高各茎蘖光合同化物生产和分配能力, 促进了分蘖发生和成穗, 获得较高的籽粒产量和水分利用效率。

关键词: 灌溉, 不同穗型小麦, 分蘖发生, 同化物生产分配, 成穗

Abstract:

Moderate irrigation based on soil moisture content could increase spike number of wheat, and realize water-saving and high-yielding cultivation, but the physiological mechanism of tiller occurrence and spike formation under this condition is still unclear. A field experiment was conducted in wheat growth seasons of 2019-2020 and 2020-2021, in which the medium-spike wheat Jimai 22 and large-spike wheat Shannong 23 were used as the experimental materials. Three treatments were designed in this field experiment, including no irrigation during the whole growth period, water-saving irrigation, and sufficient irrigation. The characteristics of tiller occurrence and spike formation of wheat were studied. The results showed that the tiller node area and trans zeatin content, the photosynthetic parameters of the uppermost unfloding leaves in main stem at wintering and turn-green stage of two wheat cultivars under water-saving irrigation treatment were higher than no irrigation treatment. Under water-saving irrigation, the stem number of tillers II, III, IP, IV, and others in two spike-type wheat cultivars were increased, the total stem number per plant of Jimai 22 and Shannong 23 under W70 were 1.01 and 0.75 higher at wintering stage and 0.71 and 0.56 higher at jointing stage. The photosynthetic parameter of the uppermost unfolding leaves at jointing stage, dry matter weight and distribution amount of 13C assimilates of main stem and tillers in different positions increased significantly, the total spike number per plant of Jimai 22 and Shannong 23 were 0.36 and 0.41 higher than no irrigation in two years. The grain yield of Jimai 22 and Shannong 23 under water-saving irrigation treatment increased by 35.00% and 44.27%, and water use efficiency increased by 9.23% and 8.55% than no irrigation, respectively. Further increasing the amount of irrigation to sufficient irrigation treatment could increase the total stem number per plant at wintering and jointing stage, but there was no significant difference with water-saving irrigation in the total formation spike number per plant, spike number, and grain yield per hectare, water use efficiency was decreased. Comparison among the cultivars the total stem and spike number per plant of Jimai 22 were higher than Shannong 23, the main reason was that Jimai 22 had higher tiller node area and trans zeatin content, photosynthetic capacity in main stem, and tillers in different positions. The correlation analysis showed that the total stem number and spike number per plant, spike number, and grain yield per hectare were positively correlated with the tiller node area and trans zeatin content, and net photosynthetic rate of the uppermost unfolding leaves in main stem in two wheat cultivars, and there was significantly negatively correlated with auxin and abscisic acid content. These results indicated that the water-saving irrigation could regulate tiller occurrence and spike formation by increasing tiller node area and hormone content, improving the capacity of photo-assimilates production and distribution in main stem and tillers in different positions, thus resulting in the higher yield and water use efficiency.

Key words: irrigation, different spike types wheat, tiller occurrence, photo-assimilates production and distribution, spike formation

图1

2019-2020和2020-2021年冬小麦生长季月平均降水量和月平均气温"

表1

灌溉对小麦越冬期不同茎蘖分蘖节面积和总面积的影响(2020-2021年)"

品种
Cultivar		 处理
Treatment		 分蘖节面积 Tiller node area (mm2) 分蘖节总面积
Total area of tiller node (mm2)
O I II 其余分蘖Other tillers
J22 W0 4.20 e 1.05 e 0.64 e 0.22 e 6.11 d
W70 6.09 b 1.20 d 1.11 c 1.80 b 10.20 b
W90 6.50 a 1.36 c 1.26 b 2.27 a 11.39 a
S23 W0 3.56 f 1.26 d 0.96 d — 5.78 e
W70 4.68 d 1.76 b 1.23 b 1.01 d 8.68 c
W90 5.06 c 1.89 a 1.36 a 1.66 c 9.97 b

图2

灌溉对小麦分蘖节激素含量和比值的影响 不同小写字母表示数值间在0.05概率水平差异显著。tZ: 反式玉米素; IAA: 生长素; ABA: 脱落酸。"

图3

灌溉对小麦越冬期和返青期主茎净光合速率、蒸腾速率和气孔导度的影响 不同小写字母表示数值间在0.05概率水平差异显著。"

图4

灌溉对小麦拔节期不同茎蘖净光合速率、蒸腾速率和气孔导度的影响 不同小写字母表示数值间在0.05概率水平差异显著。"

图5

灌溉对小麦开花期主茎净光合速率、蒸腾速率和气孔导度的影响 不同小写字母表示数值间在0.05概率水平差异显著。"

表2

灌溉对小麦不同茎蘖干物质重和13C同化物分配量的影响"

品种
Cultivar		 处理
Treatment		 干物质重
Dry matter weight (g)		 13C同化物分配量
Distribution amount of 13C assimilates (mg)
O I II III O I II III
标记后7 d (单茎) 7 days after labeling (stem)
J22 W0 1.84 d 1.64 c 1.04 c 0.57 c 2.98 b 2.52 c 1.57 c 0.43 b
W70 2.04 c 1.87 b 1.36 a 1.14 a 3.40 a 3.23 a 2.76 a 1.62 a
S23 W0 2.18 b 1.91 b 0.71 d 0.46 d 2.65 c 2.25 d 0.72 d 0.23 d
W70 2.49 a 2.37 a 1.29 b 0.61 b 3.26 a 2.92 b 2.23 b 0.34 c
成熟期(籽粒) Maturity (grain)
J22 W0 1.34 d 1.23 d 0.87 c — 0.98 b 0.94 c 0.86 c —
W70 1.50 c 1.49 c 1.19 b 0.73 a 1.36 a 1.32 a 1.16 a 0.90 a
S23 W0 1.97 b 1.59 b — — 0.88 c 0.79 d — —
W70 2.13 a 2.06 a 1.61 a — 1.32 a 1.24 b 1.09 b —
成熟期(营养器官) Maturity (vegetative organ)
J22 W0 1.41 d 1.37 c 1.19 c — 0.53 b 0.49 b 0.40 b —
W70 1.81 c 1.86 b 1.64 b 1.47 a 0.65 a 0.61 a 0.50 a 0.49 a
S23 W0 2.27 b 2.04 b — — 0.39 c 0.33 c — —
W70 2.65 a 2.62 a 2.37 a — 0.54 b 0.48 b 0.36 c —

表3

灌溉对小麦主茎叶龄、单株不同茎蘖发生数和单株总茎蘖数的影响"

生长季和
生育时期
Growth
season and growth stage		 品种
Cultivar		 处理
Treatment		 主茎叶龄
Leaf age of mean stem		 不同茎蘖发生数
Stem number in main stem and tillers in different positions per plant (stem plant-1)		 单株总茎蘖数
Total stem number
per plant
(stem plant-1)
O I II III IP IV 其余分蘖
Other
tillers
2019-2020
越冬期
Wintering		 J22 W0 5.5 c 1.00 a 1.00 a 0.83 b 0.57 d 0.34 d 0.16 c — 3.90 e
W70 6.3 b 1.00 a 1.00 a 1.00 a 0.84 a 0.62 b 0.27 b — 4.73 b
W90 6.8 a 1.00 a 1.00 a 1.00 a 0.88 a 0.73 a 0.41 a — 5.02 a
S23 W0 5.1 d 1.00 a 1.00 a 0.67 c 0.44 e 0.13 f — — 3.24 f
W70 5.7 c 1.00 a 1.00 a 0.98 a 0.64 c 0.29 e 0.17 c — 4.08 d
W90 6.1 b 1.00 a 1.00 a 0.94 a 0.73 b 0.43 c 0.24 b — 4.34 c
2020-2021
越冬期
Wintering		 J22 W0 5.3 d 1.00 a 1.00 a 0.72 c 0.30 c 0.12 d 0.07 d — 3.21 d
W70 6.1 b 1.00 a 1.00 a 1.00 a 0.71 b 0.47 b 0.22 b — 4.40 b
W90 6.5 a 1.00 a 1.00 a 0.99 a 0.84 a 0.67 a 0.30 a — 4.80 a
S23 W0 4.9 e 1.00 a 1.00 a 0.58 d 0.12 d — — — 2.70 e
W70 5.4 d 1.00 a 1.00 a 0.77 c 0.39 c 0.10 d 0.09 d — 3.35 d
W90 5.8 c 1.00 a 1.00 a 0.90 b 0.63 b 0.31 c 0.14 c — 3.98 c
2020-2021
拔节期
Jointing		 J22 W0 9.3 c 1.00 a 1.00 a 0.90 b 0.77 c 0.63 c 0.57 c 1.68 c 6.54 c
W70 10.2 b 1.00 a 1.00 a 1.00 a 0.87 b 0.76 b 0.69 b 1.93 b 7.25 b
W90 10.9 a 1.00 a 1.00 a 1.00 a 0.99 a 0.88 a 0.79 a 2.05 a 7.71 a
S23 W0 8.7 d 1.00 a 1.00 b 0.83 c 0.70 d 0.40 d 0.30 d 0.34 f 4.58 f
W70 9.5 c 1.00 a 1.00 a 0.91 b 0.80 c 0.57 c 0.43 c 0.43 e 5.14 e
W90 10.1 b 1.00 a 1.00 a 0.99 a 0.99 a 0.83 a 0.68 b 0.69 d 6.18 d

表4

灌溉对小麦成熟期不同茎蘖成穗数和单株成穗数的影响"

生长季
Growth season		 品种
Cultivar		 处理
Treatment		 不同茎蘖成穗数
Formation spike number of main stem and tillers in different positions per plant (stem plant-1)		 单株成穗数
Total Formation spike number per plant (stem plant-1)
O I II III
2019-2020 J22 W0 1.00 a 0.90 b 0.34 b — 2.24 b
W70 1.00 a 1.00 a 0.53 a 0.04 b 2.58 a
W90 1.00 a 1.00 a 0.33 b 0.20 a 2.53 a
S23 W0 1.00 a 0.42 e — — 1.42 d
W70 1.00 a 0.64 c 0.14 d — 1.79 c
W90 1.00 a 0.56 d 0.23 c — 1.80 c
2020-2021 J22 W0 1.00 a 1.00 a 0.30 c — 2.30 b
W70 1.00 a 1.00 a 0.57 a 0.11 b 2.68 a
W90 1.00 a 1.00 a 0.40 b 0.29 a 2.69 a
S23 W0 1.00 a 0.41 d — — 1.41 d
W70 1.00 a 0.67 b 0.19 d — 1.86 c
W90 1.00 a 0.60 c 0.29 c — 1.90 c
F值
F-value		 生长季 Growth season (G) — 8.33** 7.00* 16.33** 25.55**
品种 Cultivar (C) — 2045.37** 761.29** 280.33** 2448.32**
灌溉 Irrigation (I) — 78.04** 148.61** 126.33** 242.40**
G×C — 0.33ns 0.57ns 16.33** 2.89ns
G×I — 0.78ns 6.64** 4.33* 3.17ns
C×I — 30.70** 73.96** 126.33** 2.47ns
G×C×I — 6.78** 0.68ns 4.33* 0.12ns

图6

灌溉对小麦各生育时期群体总茎蘖数的影响"

表5

灌溉对小麦籽粒产量及其构成因素、总耗水量和水分利用效率的影响"

生长季
Growth season		 品种
Cultivar		 处理
Treatment		 公顷穗数
Spike number per hectare
(×104 hm-2)		 穗粒数
Grain
number per spike		 千粒重
1000-grain weight (g)		 籽粒产量
Grain yield
(kg hm-2)		 总耗水量
Water
consumption amount (mm)		 水分利用效率
Water use
efficiency
(kg hm-2 mm−1)
2019-2020 J22 W0 503.67 b 28.32 d 38.11 c 5435.99 c 326.49 e 16.64 e
W70 580.17 a 31.68 c 41.88 b 7623.94 b 419.07 c 18.19 c
W90 574.33 a 31.55 c 41.64 b 7545.21 b 471.73 a 15.99 f
S23 W0 321.33 d 42.43 b 41.33 b 5634.95 c 300.04 f 18.79 b
W70 401.83 c 46.32 a 43.20 a 8040.71 a 392.03 d 20.51 a
W90 406.33 c 45.95 a 42.58 a 7950.05 a 449.42 b 17.69 d
2020-2021 J22 W0 516.82 b 31.33 d 40.19 d 6507.55 c 373.36 e 17.43 d
W70 601.21 a 33.02 c 42.53 c 8443.04 b 443.17 c 19.05 b
W90 604.12 a 32.95 c 42.29 c 8418.14 b 516.34 a 16.30 e
S23 W0 320.15 d 42.23 b 44.33 b 5989.29 d 307.28 f 19.49 b
W70 422.58 c 45.00 a 45.96 b 8739.80 a 415.39 d 21.04 a
W90 435.76 c 44.82 a 45.27 a 8841.58 a 481.15 b 18.37 c
F值
F-value
 生长季 Growth season (G) 30.47** 4.91* 169.90** 194.76** 135.24** 37.64**
品种 Cultivar (C) 2746.76** 3101.84** 306.04ns 13.26** 179.33** 542.44**
灌溉 Irrigation (I) 295.00** 65.77** 83.53** 748.05** 1211.92** 279.32**
G×C 0.54ns 36.07** 24.68** 5.96* 12.12** 0.04ns
G×I 4.09* 3.68* 2.74ns 0.80ns 2.93ns 0.51ns
C×I 3.26ns 1.09ns 10.38ns 10.49** 5.68* 0.62ns
G×C×I 0.47ns 0.21ns 1.14ns 4.08* 5.07* 0.96ns

表6

皮尔逊相关性分析"

性状Trait TAD tZ IAA ABA NPW NPT TNJ SNP HSN GY
TAD 0.967** -0.979** -0.856** 0.954** 0.983** 0.948** 0.945** 0.966** 0.969**
tZ 0.985** -0.967** -0.908** 0.949** 0.953** 0.981** 0.864** 0.887** 0.877**
IAA -0.922** -0.913** 0.876** -0.921** -0.962** -0.941** -0.926** -0.920** -0.923**
ABA -0.939** -0.945** 0.919** -0.812** -0.804** -0.866** -0.693* -0.705* -0.744*
NPW 0.897** 0.874** -0.961** -0.935** 0.983** 0.929** 0.836** 0.912** 0.888**
NPT 0.968** 0.974* -0.833** -0.917** 0.823** 0.929** 0.913** 0.961** 0.942**
TNJ 0.919** 0.881* -0.922** -0.943** 0.919** 0.850** 0.867** 0.868** 0.869**
SNP 0.968** 0.983** -0.854** -0.913** 0.828** 0.976** 0.820** 0.970** 0.977**
HSN 0.957** 0.962** -0.806** -0.884** 0.788** 0.966** 0.797* 0.989** 0.985**
GY 0.954** 0.969** -0.820** -0.856** 0.765** 0.956** 0.774* 0.984** 0.987**
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