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作物学报 ›› 2026, Vol. 52 ›› Issue (3): 839-856.doi: 10.3724/SP.J.1006.2026.53044

所属专题: 玉米:耕作栽培·生理生化

• 耕作栽培·生理生化 • 上一篇    下一篇

外源褪黑素通过协调光反应与暗反应增强玉米苗期的耐热性

李新浩1(), 邢梦柯1, 周梓惠1, 李思烨2, 任昊2, 王洪章2, 赖华江2,*()   

  1. 1山东农业大学未来技术学院, 山东泰安 271018
    2山东农业大学农学院, 山东泰安 271018
  • 收稿日期:2025-07-01 接受日期:2025-11-18 出版日期:2026-03-12 网络出版日期:2025-12-01
  • 通讯作者: *赖华江, E-mail: lhj955297@126.com
  • 作者简介:E-mail: 2369034411@qq.com
  • 基金资助:
    山东省重点研发计划项目(2022CXPT014-02);优良品种产效协同提升关键技术研究课题项目

Exogenous melatonin enhances heat tolerance of maize at the seedling stage by coordinating light and dark reactions

Li Xin-Hao1(), Xing Meng-Ke1, Zhou Zi-Hui1, Li Si-Ye2, Ren Hao2, Wang Hong-Zhang2, Lai Hua-Jiang2,*()   

  1. 1College of Future Technology, Shandong Agricultural University, Tai’an 271018, Shandong, China
    2College of Agronomy, Shandong Agricultural University, Tai’an 271018, Shandong, China
  • Received:2025-07-01 Accepted:2025-11-18 Published:2026-03-12 Published online:2025-12-01
  • Contact: *赖华江, E-mail: lhj955297@126.com
  • Supported by:
    Key Research and Development Program of Shandong Province(2022CXPT014-02);Project of Key Technologies for Synergistic Improvement of Yield and Efficiency of Elite Varieties

摘要:

近年来, 频发的极端高温天气严重制约了我国玉米生产。外源褪黑素(Met)通过提高抗氧化能力来增强作物的抗逆性, 但其对热胁迫下玉米叶片碳同化能力及原初光化学效率的影响机制鲜有报道。本研究以热敏感型玉米品种先玉335 (XY)和耐热型玉米品种登海605 (DH)为材料, 于玉米6叶期进行热胁迫(38℃/28℃)处理, 探究外源Met对光合响应特征和荧光动力学曲线的影响。结果显示, 与清水处理相比, 外源Met促进了热胁迫下玉米叶片中可溶性糖和脯氨酸的积累, 提高了植株含水量。Met还提高了超氧化物歧化酶(SOD)、过氧化物酶(POD)和过氧化氢酶(CAT)活性, 减少了氧化胁迫和丙二醛(MDA)含量。这些改善效应降低了K相(300 μs)和L相(200 μs)的相对荧光, 缓解了热胁迫对放氧复合体的损伤和电子传递阻碍, 并提高了单位面积PSII反应中心数量。XY和DH的最大光化学效率(?Po)在Met处理下分别提高29.1%和17.2%。此外, 与清水处理相比, 外源Met改善了热胁迫下玉米叶片核酮糖-1,5-二磷酸羧化酶/加氧酶(Rubisco)和磷酸烯醇式丙酮酸羧化酶(PEPC)的活性, 尤其是在敏感品种XY中。光合响应参数显示, Met处理下, XY和DH的最大光合效率(Pnmax)分别提高了38.6%和30.8%, 光合能力(Pc)分别提高了30.2%和19.2%, 并降低了光呼吸速率(Rp), 提高了光饱和点(Isat)、CO2饱和点(Cisat)和最大羧化效率(Vcmax)。研究表明, 外源Met增强了叶片的渗透调节能力和抗氧化能力, 协同改善了光反应和暗反应, 缓解了热胁迫对玉米苗期净光合速率(Pn)的影响和对植株生长的损伤。

关键词: 玉米, 高温胁迫, 褪黑素, 叶绿素a荧光动力学, 光响应曲线, CO2响应曲线

Abstract:

In recent years, frequent extreme high-temperature events have severely limited maize production in China. Exogenous melatonin (Met) has been shown to enhance crop stress tolerance by improving antioxidant capacity; however, its role in regulating carbon assimilation and primary photochemical efficiency in maize leaves under heat stress remains poorly understood. In this study, the heat-sensitive maize cultivar Xianyu 335 (XY) and heat-tolerant cultivar Denghai 605 (DH) were used to investigate the effects of exogenous Met on photosynthetic responses and chlorophyll fluorescence kinetic curves under heat stress conditions (38℃/28℃) at the six-leaf stage. Compared with water treatment, exogenous Met significantly promoted the accumulation of soluble sugars and proline in maize leaves under heat stress, enhanced plant water content, and increased the activities of superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT), thereby reducing oxidative damage and lowering malondialdehyde (MDA) levels. These improvements alleviated the rise in relative fluorescence at the K phase (300 μs) and L phase (200 μs), mitigated heat-induced damage to the oxygen-evolving complex, relieved electron transport blockages, and increased the number of PSII reaction centers per unit area. Under Met treatment, the maximum photochemical efficiency (?Po) increased by 29.1% in XY and 17.2% in DH. Furthermore, Met enhanced the activities of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) and phosphoenolpyruvate carboxylase (PEPC), especially in the heat-sensitive XY. Photosynthetic response analysis showed that under Met treatment, the maximum net photosynthetic rate (Pnmax) increased by 38.6% in XY and 30.8% in DH, while photosynthetic capacity (Pc) increased by 30.2% and 19.2%, respectively. In addition, Met reduced the photorespiration rate (Rp) and increased the light saturation point (Isat), CO2 saturation point (Cisat), and maximum carboxylation efficiency (Vcmax). Overall, these findings indicate that exogenous Met enhances osmotic adjustment and antioxidant defense in maize leaves, synergistically improves both light and dark reactions of photosynthesis, and effectively mitigates heat stress-induced damage to net photosynthesis (Pn) and early-stage plant growth.

Key words: maize, heat stress, melatonin, chlorophyll a fluorescence kinetics, light response curve, CO2 response curve

表1

荧光瞬态OJIP曲线的JIP-test相关术语的说明和计算公式"

参数和计算公式
Parameter and formulae
参数说明
Parameter illustration
Vt = (Ft-Fo)/(Fm-Fo) O相到P相之间的相对可变荧光。Ft为脉冲光诱导t时刻的荧光强度; Fo为最小荧光(所有PSII反应中心(RC)开放时, t = 0时刻); Fm为最大荧光(所有PSII反应中心关闭时)。VjVi分别表示在J相(2 ms)和I相(30 ms)的相对可变荧光
Relative variable fluorescence between phase O and phase P. Ft, fluorescence at time t after onset of actinic illumination; Fo, minimal fluorescence, when all PSII reaction centers (RC) are open (at t = 0); Fm, maximal fluorescence, when all PSII RCs are closed. Vj and Vi represent the relatively variable fluorescence in the J phase (2 ms) and I phase (30 ms), respectively
Wk = (Ft-Fo)/(Fj-Fo) O相到J相之间的相对可变荧光。Fj为2 ms时的荧光强度
Relative variable fluorescence between phase O and phase J. Fj, fluorescence at 2 ms after onset of actinic illumination
Lband = (Ft-Fo)/(Fk-Fo) O相和K相之间的相对可变荧光。Fk为300 μs时的荧光强度
Relative variable fluorescence between phase O and phase K. Fk, fluorescence at 300 μs after onset of actinic illumination
PIabs = (ϕPo/(1-ϕPo)) (ψo/(1-ψo)) (1-ABS/RC) 基于吸收光能为基础的性能指数
Performance index on absorption basis
ϕPo = TRo/ABS = [1- (Fo/Fm)] 原初光化学的最大量子产额(在t = 0时)
Maximum quantum yield of primary photochemistry (at t = 0)
Ψo = ETo/TRo = 1-Vj 将电子传递至QA-之后的电子传递链的概率(在t = 0时)
Probability that a trapped exciton moves an electron into the electron transport chain beyond QA- (at t = 0)
ϕEo = ETo/ABS = (1-Fo/Fm) ψo 用于电子传递的量子产额(在t = 0时)
Quantum yield for electron transport (at t = 0)
ϕRo = REo/ABS = TRo/ABS (1-Vi) 光系统I (PSI)末端电子受体还原的量子产额
The quantum yield of reduction of end electron acceptors of PSI
ABS/RC = Mo (1/Vj) (1/ϕPo) 单位反应中心吸收的光能。Mo, OJIP荧光诱导曲线的初始斜率
Absorption flux per RC. Mo, the initial slope of the OJIP fluorescence induction curve
TRo/RC = Mo (1/Vj) 单位反应中心捕获的用于还原QA的能量(在t = 0时)
Energy trapped per RC for QA reduction (at t = 0)
ETo/RC = Mo (1/Vj) ψo 单位反应中心捕获的用于电子传递的能量(在t = 0时)
Energy trapped per RC for electron transport (at t = 0)
DIo/RC = ABS/RC-TRo/RC 单位反应中心耗散掉的能量(在t = 0时)
Energy dissipated per RC (at t = 0)
REo/RC = ϕRo (ETo/RC) RC为基础的电子传递到PSI受体侧末端电子受体的能量(在t = 0时)
RC-based energy for electron transport to the terminal electron acceptors at the PSI acceptor side (at t = 0)
RC/CSo = ϕPo (Vj/Mo) (ABS/CSo) 单位面积有活性反应中心的数量
Density of active reaction centers

图1

高温胁迫期间的环境温度和叶片表面温度 XYM: 先玉335在热胁迫下喷施褪黑素; XYW: 先玉335在热胁迫下喷施清水; DHM: 登海605在热胁迫下喷施褪黑素; DHW: 登海605在热胁迫下喷施清水。图A中阴影部分代表标准差(SD); 图B中柱状图显示的是平均值±SD。不同字母表示处理间差异显著(P < 0.05), n = 15 (2024)或n = 20 (2025)。"

图2

外源Met对热胁迫下玉米幼苗生长状况的影响 XYM: 先玉335在热胁迫下喷施褪黑素; XYW: 先玉335在热胁迫下喷施清水; DHM: 登海605在热胁迫下喷施褪黑素; DHW: 登海605在热胁迫下喷施清水; SPAD: 相对叶绿素含量。展示的玉米幼苗形态为2025年试验中热胁迫处理结束时所拍摄; 标尺为10 cm。柱状图显示的是平均值±SD。不同字母表示处理间差异显著(P < 0.05)。"

图3

热胁迫下喷施褪黑素对玉米幼苗气体交换参数的影响 XYM: 先玉335在热胁迫下喷施褪黑素; XYW: 先玉335在热胁迫下喷施清水; DHM: 登海605在热胁迫下喷施褪黑素; DHW: 登海605在热胁迫下喷施清水; Pn: 净光合速率; Gs: 气孔导度; Ci: 胞间CO2浓度; Tr: 蒸腾速率。柱状图显示的是平均值±SD。不同字母表示处理间差异显著(P < 0.05), n = 6。"

图4

热胁迫下喷施褪黑素对玉米幼苗光合响应曲线的影响 XYM: 先玉335在热胁迫下喷施褪黑素; XYW: 先玉335在热胁迫下喷施清水; DHM: 登海605在热胁迫下喷施褪黑素; DHW: 登海605在热胁迫下喷施清水; Pn: 净光合速率; PARi: 光合有效辐射; Ci: 胞间CO2浓度。"

表2

热胁迫下喷施褪黑素对光合-光响应曲线拟合参数的影响"

年份 Year 品种
Cultivar
处理
Treatment
暗呼吸速率
Rd (μmol m-2 s-1)
最大净光合速率Pnmax (μmol m-2 s-1) 饱和光强
Isat (μmol m-2 s-1)
光补偿点
Ic (μmol m-2 s-1)
表观量子效率
AQY (mmol m-2 s-1)
2024 XY Met 2.77±0.60 a 14.96±0.60 a 1044.03±45.50 a 59.69±5.56 a 0.038±0.005 a
Water 2.58±0.41 a 11.56±0.64 b 910.61±27.79 b 68.40±12.45 a 0.032±0.006 a
2025 XY Met 3.36±0.73 a 24.00±0.47 a 1460.49±8.75 b 65.67±12.15 a 0.040±0.003 a
Water 4.04±0.50 a 17.31±1.52 c 1219.31±107.46 c 68.15±7.49 a 0.050±0.008 a
DH Met 3.53±0.42 a 19.85±0.48 b 1917.19±64.26 a 72.36±3.84 a 0.040±0.005 a
Water 3.66±0.15 a 15.18±0.43 d 1428.02±165.9 bc 73.20±12.65 a 0.040±0.003 a
方差分析
ANOVA
品种
Cultivar (C)
ns ** ** ns ns
处理Treatment (T) ns ** ** ns ns
C × T ns ns ns ns ns

表3

热胁迫下喷施褪黑素对光合-CO2响应曲线拟合参数的影响"

年份
Year
品种
Cultivar
处理
Treatment
光呼吸速率
Rp (μmol m-2 s-1)
光合能力
Pc (μmol m-2 s-1)
饱和胞间CO2浓度 Cisat (μmol mol-l) CO2补偿点
Γ (μmol mol-l)
最大羧化速率 Vcmax (μmol m-2 s-1)
2024 XY Met 14.73±2.15 b 14.86±0.54 a 558.11±37.48 a 32.73±0.48 a 17.63±0.45 a
Water 35.48±2.31 a 10.71±0.05 b 470.25±12.26 b 35.73±2.06 a 13.30±0.37 b
2025 XY Met 4.57±1.28 b 27.10±0.54 a 355.88±50.92 b 4.81±0.49 b 47.55±2.63 a
Water 9.80±0.56 a 22.27±0.45 b 404.69±53.29 b 33.13±5.14 a 35.21±0.83 b
DH Met 1.18±0.37 c 19.61±0.86 c 760.57±174.45 a 3.51±0.55 b 26.29±4.17 c
Water 1.88±0.55 c 16.45±0.45 d 481.59±62.78 b 4.11±0.38 b 21.67±1.55 c
方差分析
ANOVA
品种
Cultivar (C)
** ** ** ** **
处理
Treatment (T)
** ** ns ** **
C × T ** ns * ** ns

图5

热胁迫下喷施褪黑素对玉米幼苗叶绿素a荧光瞬态、O-P相的相对可变荧光(Vt)、处理Vt与对照Vt的差值(ΔVt)、O-J相的相对可变荧光(Wk)和O-K相的相对可变荧光(Lband)的影响 XYM: 先玉335在热胁迫下喷施褪黑素; XYW: 先玉335在热胁迫下喷施清水; DHM: 登海605在热胁迫下喷施褪黑素; DHW: 登海605在热胁迫下喷施清水; XYCK: 在正常环境下生长的先玉335; DHCK: 在正常环境下生长的登海605; Vt: O-P相的相对可变荧光强度; ΔVt: O-P相的相对可变荧光强度与对照的差值; Wk: O相到J相的相对可变荧光; ΔWk: O相到J相之间的相对可变荧光与对照的差值; Lband: O相和K相的相对可变荧光; ΔLband: O相和K相之间的相对可变荧光与对照的差值。"

图6

热胁迫下喷施褪黑素对玉米幼苗叶绿素a荧光瞬态JIP-test相关参数的影响 XYM: 先玉335在热胁迫下喷施褪黑素; XYW: 先玉335在热胁迫下喷施清水; DHM: 登海605在热胁迫下喷施褪黑素; DHW: 登海605在热胁迫下喷施清水; CK: 在正常环境下生长的先玉335或登海605; PIabs: 基于吸收光能为基础的性能指数; Vj: J相(2 ms)的相对可变荧光强度; Vi: I相(30 ms)的相对可变荧光强度; Fo: 所有PSII反应中心开放时的荧光强度; Fm: 最大荧光强度; ABS/RC: 单位反应中心吸收的光能; DIo/RC: 单位反应中心耗散掉的能量; TRo/RC: 单位反应中心捕获的用于还原QA的能量; ETo/RC: 单位反应中心捕获的用于电子传递的能量; REo/RC: RC为基础的电子传递到PSI受体侧末端电子受体的能量; ?Po: 原初光化学的最大量子产额; ψo: 将电子传递至QA-之后的电子传递链的概率; ?Eo: 用于电子传递的量子产额; ?Ro: PSI末端电子受体还原的量子产额; RC/CSo: 单位面积有活性反应中心的数量。"

图7

热胁迫下喷施褪黑素对玉米叶片抗氧化酶活性和MDA含量的影响 XYM: 先玉335在热胁迫下喷施褪黑素; XYW: 先玉335在热胁迫下喷施清水; DHM: 登海605在热胁迫下喷施褪黑素; DHW: 登海605在热胁迫下喷施清水; SOD: 超氧化物歧化酶; POD: 过氧化物酶; CAT: 过氧化氢酶; MDA: 丙二醛。单位中的A560表示560 nm波长下的吸光度, Δ240和Δ470分别表示单位时间内240 nm和470 nm波长下吸光度的变化值。柱状图显示的是平均值±标准差。不同字母表示处理间差异显著(P < 0.05), n = 3。"

图8

热胁迫下喷施褪黑素对玉米叶片渗透调节物质含量的影响 XYM: 先玉335在热胁迫下喷施褪黑素; XYW: 先玉335在热胁迫下喷施清水; DHM: 登海605在热胁迫下喷施褪黑素; DHW: 登海605在热胁迫下喷施清水。柱状图显示的是平均值±标准差。不同字母表示处理间差异显著(P < 0.05), n = 3。"

图9

热胁迫下喷施褪黑素对玉米幼苗叶片Rubisco和PEPC活性的影响 XYM: 先玉335在热胁迫下喷施褪黑素; XYW: 先玉335在热胁迫下喷施清水; DHM: 登海605在热胁迫下喷施褪黑素; DHW: 登海605在热胁迫下喷施清水; Rubisco: 核酮糖-1,5-二磷酸羧化酶/加氧酶; PEPC: 磷酸烯醇式丙酮酸羧化酶。柱状图显示的是平均值±标准差。不同字母表示处理间差异显著(P < 0.05), n = 3。"

图10

本试验条件下各指标的皮尔逊相关性分析和对地上部生物量的相对重要性分析 ADW: 地上部干重; WC: 植株含水量; SPAD: 相对叶绿素含量; Ci: 胞间CO2浓度; Gs: 气孔导度; Pn: 净光合速率; Tr: 蒸腾速率; Rd: 暗呼吸速率; Pnmax: 最大净光合速率; Isat: 饱和光强; Ic: 光补偿点; AQY: 表观量子效率; Rp: 光呼吸速率; Pc: 光合能力; Vcmax: 最大羧化速率; Cisat: 饱和胞间CO2浓度; Γ: CO2补偿点; Lband: O相和K相之间的相对可变荧光; Wk: O相到J相之间的相对可变荧光; Vj: J相(2 ms)的相对可变荧光强度; Vi: I相(30 ms)的相对可变荧光强度; PIabs: 基于吸收光能为基础的性能指数; Fo: 所有PSII反应中心开放时的荧光强度; Fm: 最大荧光强度; ABS/RC: 单位反应中心吸收的光能; DIo/RC: 单位反应中心耗散掉的能量; TRo/RC: 单位反应中心捕获的用于还原QA的能量; ETo/RC: 单位反应中心捕获的用于电子传递的能量; REo/RC: RC为基础的电子传递到PSI受体侧末端电子受体的能量; ?Po: 原初光化学的最大量子产额; ψo: 将电子传递至QA-之后的电子传递链的概率; ?Eo: 用于电子传递的量子产额; ?Ro: PSI末端电子受体还原的量子产额; RC/CSo: 单位面积有活性反应中心的数量; SOD: 超氧化物歧化酶活性; POD: 过氧化物酶活性; CAT: 过氧化氢酶活性; MDA: 丙二醛含量; SS: 可溶性糖含量; SP: 可溶性蛋白质含量; Pro: 脯氨酸含量; Rubisco: 核酮糖-1,5-二磷酸羧化酶/加氧酶; PEPC: 磷酸烯醇式丙酮酸羧化酶。"

图11

热胁迫下外源褪黑素改善玉米幼苗光合性能和植株生长的作用机制 Met: 褪黑素; WC: 植株含水量; Pn: 净光合速率; Isat: 饱和光强; Ic: 光补偿点; Rp: 光呼吸速率; Pc: 光合能力; Vcmax: 最大羧化速率; ΔLband: O相和K相之间的相对可变荧光与对照的差值; ΔWk: O相到J相之间的相对可变荧光与对照的差值; Vk: K相(300 μs)的相对可变荧光强度; Vi: I相(30 ms)的相对可变荧光强度; PIabs: 基于吸收光能为基础的性能指数; ?Po: 原初光化学的最大量子产额; ?Eo: 用于电子传递的量子产额; RC/CSo: 单位面积有活性反应中心的数量; SOD: 超氧化物歧化酶活性; POD: 过氧化物酶活性; CAT: 过氧化氢酶活性; MDA: 丙二醛含量; SS: 可溶性糖含量; SP: 可溶性蛋白质含量; Pro: 脯氨酸含量; Rubisco: 核酮糖-1,5-二磷酸羧化酶/加氧酶; PEPC: 磷酸烯醇式丙酮酸羧化酶。"

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