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作物学报 ›› 2022, Vol. 48 ›› Issue (11): 2920-2933.doi: 10.3724/SP.J.1006.2022.11109

• 耕作栽培·生理生化 • 上一篇    下一篇

土壤水分亏缺和大气CO2浓度升高对冬小麦光合特性的影响

郑云普1(), 常志杰1, 韩怡1, 卢云泽2, 陈文娜2, 田银帅2, 殷嘉伟2, 郝立华1,3,*()   

  1. 1河北工程大学水利水电学院, 河北邯郸 056038
    2河北工程大学园林与生态工程学院, 河北邯郸 056038
    3南京农业大学园艺学院, 江苏南京 210095
  • 收稿日期:2021-12-03 接受日期:2022-03-25 出版日期:2022-11-12 网络出版日期:2022-04-21
  • 通讯作者: 郝立华
  • 作者简介:第一作者联系方式: E-mail: zhengyunpu@hebeu.edu.cn
  • 基金资助:
    本研究由国家自然科学基金项目(32071608);河北省自然科学基金项目(C2020402026);河北省自然科学基金项目(E2021402031)

Effects of soil water deficit and elevated atmospheric CO2 concentration on leaf photosynthesis of winter wheat

ZHENG Yun-Pu1(), CHANG Zhi-Jie1, HAN Yi1, LU Yun-Ze2, CHEN Wen-Na2, TIAN Yin-Shuai2, YIN Jia-Wei2, HAO Li-Hua1,3,*()   

  1. 1School of Water Conservancy and Hydropower, Hebei University of Engineering, Handan 056038, Hebei, China
    2School of Landscape and Ecological Engineering, Hebei University of Engineering, Handan 056038, Hebei, China
    3College of Horticulture, Nanjing Agricultural University, Nanjing 210095, Jiangsu, China
  • Received:2021-12-03 Accepted:2022-03-25 Published:2022-11-12 Published online:2022-04-21
  • Contact: HAO Li-Hua
  • Supported by:
    The National Natural Science Foundation of China(32071608);The Natural Science Foundation of Hebei Province(C2020402026);The Natural Science Foundation of Hebei Province(E2021402031)

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摘要：

为深入理解未来大气CO2浓度([CO2])升高背景下农田生态系统结构与功能对土壤水分亏缺的响应机制, 利用可精准控制[CO2]的大型环境生长箱, 研究了土壤水分亏缺和大气[CO2]升高对冬小麦气孔特征、净光合速率、水分利用效率、叶片碳氮含量、非结构性碳水化合物含量、核酮糖-1,5-二磷酸羧化酶/加氧酶(Rubisco)活性及其基因表达量的影响。本研究结果表明, 水分亏缺导致冬小麦的总生物量和净光合速率(Pn)分别相比对照降低33%和29%, 而[CO2]升高可以在一定程度上缓解水分亏缺对冬小麦生长及生理过程造成的不利影响。同时, 水分亏缺还使冬小麦气孔开度及其空间分布格局规则性的降低, 但高[CO2]可以通过增加气孔密度和提高气孔分布的规则程度, 进一步优化冬小麦叶片的气体交换效率。另外, [CO2]升高增加水分亏缺条件下冬小麦的Pn, 但同时却导致蒸腾速率(Tr)降低25%, 从而提高叶片的瞬时水分利用效率(WUEI)61%。此外, 水分亏缺条件下[CO2]升高不仅导致Rubisco酶初始活性、活化率以及可溶性蛋白含量分别增加66%、38%和15%, 而且还分别提高Rubisco酶编码基因RbcL3和RbcS2的表达水平453%和417%。上述结果表明, 水分亏缺条件下, [CO2]升高可以通过调整气孔特征和Rubisco酶活性及其编码基因表达水平, 进一步优化冬小麦的气体交换效率, 从而提高植株生物量、净光合速率以及水分利用效率。研究结果将为深入理解未来气候变化背景下冬小麦响应[CO2]升高和水分亏缺的生理及分子机制提供理论依据。

关键词: 土壤水分亏缺, CO2浓度升高, 冬小麦, 气孔特征, 光合性能, Rubisco基因表达

Abstract:

To understand the mechanisms of agricultural ecosystem structure and function in response to soil water deficit under future elevated atmospheric CO2 concentration, we examined the effects of soil water deficit and elevated CO2 concentration ([CO2]) on the stomatal traits, leaf photosynthesis, water use efficiency as well as Rubisco activity and gene expression of winter wheat with environmental growth chambers, whereby the [CO2] was controlled accurately with CO2 sensors. Our results showed that water deficit significantly decreased the plant biomass and net photosynthesis rates (Pn) of winter wheat by 33% and 29%, whereas elevated [CO2] partially mitigated the negative effects of water deficit on plant growth and physiological processes of winter wheat. Meanwhile, water deficit also reduced the stomatal width and regularity of stomatal distribution pattern on winter wheat leaves, but higher [CO2] could optimize the leaf gas exchange efficiency with more regular distribution pattern of stomata. Moreover, elevated [CO2] not only enhanced the Pn of winter wheat plants under water deficit, but also substantially reduced the transpiration rates (Tr) by 25%, and thus elevated [CO2] increased the water use efficiency by 61% when winter wheat plants subjected to water deficit. In addition, elevated [CO2] boosted the initial activity and activation state of Rubisco as well as soluble protein content by 66%, 38%, and 15%, and meanwhile significantly enhanced the gene expression levels of RbcL3 and RbcS2 by 453% and 417%, respectively. These results suggested that elevated [CO2] may optimize leaf gas exchange through modifying stomatal traits as well as the activity and gene express of Rubisco, and thus increased plant biomass, Pn, and water use efficiency to efficiently alleviate the physiological stress of water deficit on growth and development processes of winter wheat. Our findings may not only provide data for further understanding the impacts of water deficit on grain yield and water use efficiency of winter wheat under elevated [CO2], but also have important significance for adaptation management of agricultural ecosystems under global change.

Key words: water deficiency, doubling CO2 concentration, winter wheat, stomatal traits, photosynthetic performance, Rubisco gene expression

图1

水分亏缺和[CO2]升高对冬小麦叶面积(a)及生物量(b)的影响 不同字母标注表示处理间为显著性差异(P < 0.05), 相同字母标注表示处理间为不显著性差异(P > 0.05)。"

表1

水分亏缺和[CO2]升高对冬小麦叶面积和生物量的交互影响"

处理因素 Treatment 叶面积 Leaf area 植株生物量 Plant biomass
CO2浓度 CO2 concentration ([CO2]) 0.371 0.117
水分 Watering <0.001 <0.001
[CO2]×水分 [CO2] × Watering 0.005 0.199

表2

水分亏缺和[CO2]升高对冬小麦气孔形态参数的影响"

气孔形态特征
Stomatal morphology		 叶面
Leaf surface		 对照
Control		 水分亏缺
Water deficit		 [CO2]升高
e[CO2]		 水分亏缺×[CO2]升高
Water deficit × e[CO2]		 P值
P-value
气孔密度
Stomatal density		 近轴面Adaxial 51.1±1.2 b 61.1±2.0 a 64.8±2.6 a 60.1±2.3 a <0.001
远轴面Abaxial 29.3±1.2 c 32.5±0.8 bc 40.3±2.3 a 34.7±0.8 b <0.001
气孔长度
Stomatal length (μm)		 近轴面Adaxial 40.4±1.2 a 38.9±2.0 a 38.0±0.5 a 40.3±1.4 a 0.402
远轴面Abaxial 37.0±1.1 b 38.9±1.0 ab 39.4±1.6 ab 42.5±0.9 a 0.076
气孔宽度
Stomatal width (μm)		 近轴面Adaxial 4.1±0.2 b 4.8±0.2 a 3.6±0.1 c 3.5±0.1 c <0.001
远轴面Abaxial 4.2±1.5 a 4.3±0.4 a 4.2±1.0 a 3.5±0.3 a 0.338
气孔周长
Stomatal perimeter (μm)		 近轴面Adaxial 85.2±2.5 a 82.5±3.8 a 79.3±1.2 a 83.9±2.8 a 0.337
远轴面Abaxial 78.3±2.5 b 82.3±2.0 ab 83.4±3.6 ab 88.8±1.7 a 0.163
气孔面积
Stomatal area (μm2)		 近轴面Adaxial 167.4±15.6 a 191.4±13.0 a 129.5±6.1 b 129.1±6.2 b 0.002
远轴面Abaxial 146.4±20.0 a 167.9±8.0 a 173.7±18.2 a 146.9±3.6 a 0.521
气孔形状指数
Stomatal shape index (%)		 近轴面Adaxial
远轴面Abaxial		 0.15±0.012 b 0.17±0.003 a 0.14±0.007 bc 0.14±0.009 c <0.001
0.15±0.006 a 0.16±0.002 a 0.15±0.003 a 0.14±0.003 b 0.011

表3

水分和[CO2]对冬小麦气孔形态特征的交互影响"

气孔形态特征
Stomatal morphology		 气孔密度
Stomatal density		 气孔长度
Stomatal length		 气孔宽度
Stomatal width		 气孔周长
Stomatal perimeter		 气孔面积
Stomatal area		 形状指数
Shape
index
CO2浓度 CO2 concentration ([CO2]) <0.001 0.198 0.001 0.377 0.028 <0.001
水分 Watering 0.547 0.118 0.715 0.164 0.662 0.866
叶面 Leaf surface <0.001 0.984 0.904 0.816 0.677 0.964
[CO2]×水分 [CO2] × Watering <0.001 0.184 0.019 0.288 0.086 <0.001
[CO2]×叶面 [CO2] × Leaf surface 0.909 0.061 0.065 0.049 0.013 0.042
水分×叶面 Watering × Leaf surface 0.107 0.255 0.178 0.354 0.489 0.054
[CO2]×水分×叶面 [CO2] × Watering × Leaf surface 0.230 0.479 0.741 0.466 0.568 0.984

图2

水分亏缺和[CO2]升高对冬小麦近轴面(a)和远轴面(b)气孔分布格局的影响 图中的Upper 95%表示95%置信区间上界线, Lower 95%表示95%置信区间下界线。CK代表Control,D代表干旱处理,ED代表CO2浓度升高和干旱处理,E代表CO2浓度升高处理。"

图3

水分亏缺和[CO2]升高对冬小麦叶片气体交换参数的影响 不同字母标注表示处理间为显著性差异(P < 0.05), 相同字母标注表示处理间为不显著性差异(P > 0.05)。"

表4

水分亏缺和[CO2]升高对冬小麦叶片气体交换的交互影响"

气体交换
Gas exchange		 净光合速率
Pn		 蒸腾速率
Tr		 气孔导度
Gs		 瞬时水分利用率
WUEI		 暗呼吸速率
Rd		 细胞间[CO2]
Ci
CO2浓度 CO2 concentration ([CO2]) 0.071 0.143 0.044 <0.001 0.211 0.278
水分 Watering 0.008 <0.001 0.002 <0.001 <0.001 <0.001
[CO2]×水分 [CO2] × Watering 0.369 0.071 0.019 0.567 0.275 0.144

表5

水分亏缺和[CO2]升高对冬小麦叶片非结构性碳水化合物的交互影响"

处理因素
Treatment		 果糖
Fructose		 蔗糖
Sucrose		 葡萄糖
Glucose		 可溶性糖
Soluble sugars		 淀粉
Starch		 非结构性碳水化合物
Total NSC
CO2浓度 CO2 concentration ([CO2]) 0.001 <0.001 0.098 0.256 0.293 0.499
水分 Watering 0.352 0.098 0.001 0.877 0.618 0.526
[CO2]×水分 [CO2] × Watering 0.001 <0.001 0.001 <0.001 0.028 0.944

图4

水分亏缺和[CO2]升高对冬小麦叶片非结构性碳水化合物的影响 不同字母标注表示处理间为显著性差异(P < 0.05), 相同字母标注表示处理间为不显著性差异(P > 0.05)。"

表6

水分亏缺和[CO2]升高对冬小麦叶片C、N和C/N的交互影响"

处理因素
Treatment		 C含量
Carbon content		 N含量
Nitrogen content		 C/N
C/N ratio
CO2浓度 CO2 concentration ([CO2]) 0.004 0.055 0.151
水分 Watering 0.308 0.841 0.904
[CO2]×水分 [CO2] × Watering 0.443 0.293 0.438

图5

水分亏缺和[CO2]升高对冬小麦叶片C、N含量及C/N的影响 不同字母标注表示处理间为显著性差异(P < 0.05), 相同字母标注表示处理间为不显著性差异(P > 0.05)。"

图6

水分亏缺和[CO2]升高对冬小麦叶片Rubisco活性和可溶性蛋白的影响"

图7

水分亏缺和[CO2]升高对冬小麦叶片基因表达量的影响 不同字母标注表示处理间为显著性差异(P < 0.05), 相同字母标注表示处理间为不显著性差异(P > 0.05)。"

图8

水分亏缺和[CO2]升高协同影响冬小麦叶片光合、蒸腾及水分利用的过程及其潜在机制[19]"

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