作物杂志, 2026, 42(2): 154-159 doi: 10.16035/j.issn.1001-7283.2026.02.019

生理生化·植物营养·栽培耕作

外源黄体酮影响小麦根系伸长生长的生理调控途径

王丹,1, 王润2, 胡孝庆2, 于会勇1, 李江涛1, 程星1, 郭海悦1, 刘婷1, 胡珍珍1, 李华,2

1濮阳市农林科学院457000河南濮阳

2河南农业大学生命科学学院450002河南郑州

Physiological Regulation Pathways of Exogenous Progesterone Affecting Wheat Root Elongation and Growth

Wang Dan,1, Wang Run2, Hu Xiaoqing2, Yu Huiyong1, Li Jiangtao1, Cheng Xing1, Guo Haiyue1, Liu Ting1, Hu Zhenzhen1, Li Hua,2

1Puyang Academy of Agricultural and Forestry Sciences, Puyang 457000, Henan, China

2College of Life Sciences, Henan Agricultural University, Zhengzhou 450002, Henan

通讯作者: 李华,主要从事作物逆境生理与分子调控机制研究,E-mail:lihua@henau.edu.cn

收稿日期: 2025-01-7   修回日期: 2025-02-18   网络出版日期: 2025-05-08

基金资助: 河南省科技攻关项目(242102111147)
河南省重大科技专项(241100110100)
濮阳市重大科技攻关项目(230119)
财政部和农业农村部:国家现代农业产业技术体系(CARS-03-72)

Received: 2025-01-7   Revised: 2025-02-18   Online: 2025-05-08

作者简介 About authors

王丹,主要从事农作物新品种选育与栽培技术研究,E-mail:pywangdan2012@sina.cn

摘要

黄体酮是一种存在于植物体中的类固醇激素,与植物的生长发育紧密相关,对植物抵抗逆境胁迫也有积极作用。小麦根系生长状况直接影响产量与抗逆能力,本试验研究了外源黄体酮对小麦根系伸长生长的影响,并初步分析其调控途径。结果表明,不同浓度外源黄体酮对小麦根系生长的调控效应存在差异,低浓度(0.001和0.01 μmol/L)促进根系伸长生长,高浓度(0.1和1.0 μmol/L)抑制其生长。外源黄体酮处理显著影响小麦幼苗根系中的葡萄糖含量及磷酸果糖激酶(PFK)活性,随着黄体酮处理浓度升高,根系葡萄糖含量先降后升,而PFK活性变化趋势与之相反。进一步研究发现,10 μmol/L葡萄糖处理下幼苗根系葡萄糖含量降低,PFK活性升高;10 000 μmol/L葡萄糖处理下幼苗根系葡萄糖含量升高,PFK活性受到抑制。此外,添加0.1 μmol/L黄体酮可显著促进低浓度葡萄糖处理下根系葡萄糖的积累,并抑制葡萄糖对PFK活性的诱导作用;而添加0.001 μmol/L黄体酮能显著抑制高浓度葡萄糖处理下根系葡萄糖的累积,缓解葡萄糖对PFK活性的抑制作用。外源黄体酮可能通过作用于糖酵解途径的关键限速酶PFK影响糖酵解过程,进而调控根系中葡萄糖含量,实现对小麦根系伸长生长的调控。

关键词: 小麦; ; 黄体酮; 葡萄糖; 磷酸果糖激酶

Abstract

Progesterone is a steroid hormone found in plants, which is closely related to plant growth and development and plays a positive role in stress resistance. The growth status of wheat roots directly affects yield and stress resistance. This study investigated the effects of exogenous progesterone on the elongation growth of wheat roots and preliminarily analyzed their regulatory pathways. The results showed that different concentrations of exogenous progesterone exerted varying regulatory effects on wheat root growth: low concentrations (0.001 and 0.01 μmol/L) promoted root elongation, whereas high concentrations (0.1 and 1.0 μmol/L) inhibited growth. Exogenous progesterone treatment significantly affected the glucose content and phosphofructokinase (PFK) activity in the roots of wheat seedlings. With increasing progesterone concentration, the root glucose content first decreased and then increased, while the trend of PFK activity was opposite. Further research found that under 10 μmol/L glucose treatment, the root glucose content decreased and PFK activity increased; under 10 000 μmol/L glucose treatment, the root glucose content increased and PFK activity was inhibited. In addition, the application of 0.1 μmol/L progesterone significantly promoted root glucose accumulation under low-concentration glucose treatment and inhibited the induction effect of glucose on PFK activity. Conversely, the application of 0.001 μmol/L progesterone significantly inhibited root glucose accumulation under high-concentration glucose treatment and alleviated the inhibitory effect of glucose on PFK activity. These findings suggest that exogenous progesterone may affect the glycolysis process by acting on PFK, a key rate-limiting enzyme in the glycolytic pathway, thereby regulating the glucose content in roots and regulating wheat root elongation growth.

Keywords: Wheat; Root; Progesterone; Glucose; Phosphofructokinase

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本文引用格式

王丹, 王润, 胡孝庆, 于会勇, 李江涛, 程星, 郭海悦, 刘婷, 胡珍珍, 李华. 外源黄体酮影响小麦根系伸长生长的生理调控途径. 作物杂志, 2026, 42(2): 154-159 doi:10.16035/j.issn.1001-7283.2026.02.019

Wang Dan, Wang Run, Hu Xiaoqing, Yu Huiyong, Li Jiangtao, Cheng Xing, Guo Haiyue, Liu Ting, Hu Zhenzhen, Li Hua. Physiological Regulation Pathways of Exogenous Progesterone Affecting Wheat Root Elongation and Growth. Crops, 2026, 42(2): 154-159 doi:10.16035/j.issn.1001-7283.2026.02.019

黄体酮是普遍存在于高等植物中的一种激素,在不同植物各器官中的含量差异较大[1-2],功效也不尽相同。近年来研究表明,黄体酮在多种逆境胁迫中发挥重要作用,适宜浓度的黄体酮可增强植物体内抗氧化酶活性,降低脂质过氧化水平和过氧化氢含量,缓解盐胁迫对小麦幼苗生长的抑制作用[3],改变低温胁迫下小麦的细胞膜脂结构[4],还可减轻高温和高光强引发的氧化伤害[5-6]。在拟南芥中,黄体酮预处理可缓解假单胞菌侵染导致的坏死现象[7]。过表达CYP11A1的烟草转基因植株(黄体酮水平比野生型植株高3~4倍)能有效抵御真菌病原体侵染[8]。此外,外施黄体酮可减轻鹰嘴豆、玉米及香蕉果实在冷胁迫下诱导的氧化伤害[9-11]

黄体酮不仅在植物抵御逆境胁迫方面发挥积极作用,还与植物的生长发育紧密相关。研究[1]表明,低浓度黄体酮(0.01~1.00 μmol/L)可促进拟南芥幼苗生长,而高浓度(100 μmol/L)则对其生长产生抑制作用。高浓度黄体酮(0.25 μg/株)能促进向日葵地上部分生长,低浓度(0.1 μg/株)则有利于其根的伸长[12]。此外,黄体酮可介导植物花的发育过程。外源施加黄体酮能够促进烟草花粉管生长[13];猕猴桃内源黄体酮含量随花粉萌发逐渐升高[14]。在小麦[15-16]和拟南芥[17]中,黄体酮还被发现具有诱导开花和促进生殖生长的作用。

黄体酮虽能调节植物生长发育,但其具体调控机制相关的研究尚显不足。目前,黄体酮相关研究多集中于洋地黄、苦参、桂竹香和小球藻等植物[18],在粮食作物中的研究相对较少,针对植物根系的研究则更为匮乏。小麦(Triticum aestivum L.)是全球主要粮食作物之一,研究黄体酮对其生长发育的影响,对小麦抗逆性研究及生产实践具有重要意义。根系在小麦生长中起到固定植株、提供养分和水分等重要作用,直接影响其产量形成及植株抗逆能力。本试验以小麦为研究对象,重点探究黄体酮对小麦根系伸长生长的影响,并解析其调控机制及主要途径,旨在为揭示黄体酮调控作物根系生长的路径以及指导小麦抗逆稳产栽培提供理论依据。

1 材料与方法

1.1 试验设计

以黄淮地区主推小麦品种“百农207”为试验材料。选取籽粒饱满且大小均匀的种子,用5%过氧化氢消毒5 min,蒸馏水冲洗3~4次。用蒸馏水浸泡4~6 h后,把种子放在湿润的滤纸上催芽72 h,待种子芽长约2 cm、主根长约3 cm时,将长势一致的幼苗移到不同浓度外源黄体酮和葡萄糖的Hoagland培养液中进行培养,环境条件为:温度25 oC/22 oC(白天/晚上),光周期14 h/10 h(光照/黑暗),光照强度300 μmol/(m2·s),相对湿度保持在70%。

设5种浓度的黄体酮处理,分别为0.000(P0)、0.001(P1)、0.010(P2)、0.100(P3)和1.000 μmol/L(P4);7种浓度的葡萄糖Hoagland培养液处理,分别为0.0(G0)、0.1(G1)、1.0(G2)、10.0(G3)、100.0(G4)、1000.0(G5)和10 000.0 μmol/L(G6)。黄体酮和葡萄糖交互处理试验共设置5个处理组合,分别为正常培养(PG0,CK)、10 μmol/L葡萄糖(PG1)、10 μmol/L葡萄糖+0.1 μmol/L黄体酮(PG2)、10 000 μmol/L葡萄糖(PG3)和10 000 μmol/L葡萄糖+0.001 μmol/L黄体酮(PG4)。每个处理不少于3次重复,每3 d更换1次营养液,6 d后测定根长,并取样保存于-80 oC超低温冰箱备测,各处理样品测试3次重复。

1.2 测定项目与方法

1.2.1 根长

在根系旁放置标尺并拍照,随后将图片置入Photoshop 7.0,使用标尺工具测定根长,每种处理测量5~10株,结果取平均值。

1.2.2 葡萄糖含量

将0.1 g样品于预冷研钵中迅速研磨,转移至1.5 mL的离心管中,加入1 mL蒸馏水,置沸水浴中煮沸10 min。室温冷却后,8000 g室温离心10 min,取上清液,采用BC2500试剂盒(北京索莱宝科技有限公司)测定葡萄糖含量。

1.2.3 磷酸果糖激酶(PFK)活性

将0.1 g样品于预冷研钵中迅速研磨,转移至1.5 mL的离心管中,加入1 mL提取液,采用冷冻离心机在4 ℃下8 000 g离心10 min,取上清液,采用BC0530试剂盒(北京索莱宝科技有限公司)测定PFK活性。

1.3 数据处理

采用SPSS软件进行数据统计与分析,采用Duncan新复极差法进行差异显著性检验。

2 结果与分析

2.1 外源黄体酮对小麦根系生长的影响

图1所示,在P1和P2处理下,小麦平均根长较P0处理显著增长,分别增加了15.3%和9.0%,但P1和P2处理间差异不显著;P3和P4处理的平均根长较P0处理分别显著减少了10.9%和34.6%,2个处理间差异也达显著水平,表明低浓度外源黄体酮促进小麦根系伸长,高浓度抑制其伸长。

图1

图1   外源黄体酮对小麦根系生长的影响

不同小写字母表示处理间在P < 0.05水平差异显著,下同。

Fig.1   Effects of exogenous progesterone on root elongation of wheat

Different lowercase letters indicate significant difference among treatments at P < 0.05 level, the same below.


2.2 外源黄体酮对小麦根系中葡萄糖含量及PFK活性的影响

图2所示,与P0处理相比,P1和P2处理均降低了小麦根系中的葡萄糖含量,P1处理根系中葡萄糖含量降为P0处理的65%,与P0处理差异显著;P2处理略有降低,与P0处理差异不显著;且P1与P2处理间差异显著。P3和P4处理的根系中葡萄糖含量显著升高,分别为P0处理的1.35和2.23倍,且2个处理间的差异也达到显著水平。综上,低浓度黄体酮处理下,小麦根系中葡萄糖含量降低;高浓度黄体酮处理下,根系中葡萄糖含量显著增加。

图2

图2   外源黄体酮对小麦根系中葡萄糖含量和PFK活性的影响

Fig.2   Effects of exogenous progesterone on glucose content and PFK activity in wheat roots


PFK是糖酵解代谢途径中的关键限速酶,如图2b所示,P1和P2处理的PFK活性显著增加,较P0处理分别提高了30.4%和21.6%,2个处理间差异不显著;P3和P4处理的小麦根系中PFK活性显著下降,分别为P0处理的89.5%和49.7%,差异达显著水平,且P3和P4处理间差异显著。综上,低浓度的黄体酮诱导PFK活性增加,而高浓度黄体酮抑制PFK活性。

外源黄体酮显著影响小麦幼苗根系中的葡萄糖含量和PFK的活性。随着外源黄体酮浓度升高,小麦根系中的葡萄糖含量先降低后升高,而PFK活性则呈先升高后下降的趋势。

2.3 外源葡萄糖对小麦根系生长的影响

除G6处理外,其余处理的根长较G0处理均有不同程度增长(图3)。其中,G1、G2和G3处理与G0处理的差异达显著水平,G3处理增长最多,增幅为62.9%;G4和G5处理增长不显著;G6处理根长仅为G0处理的68.5%,差异显著。综上,外源葡萄糖可显著影响小麦根长,低浓度促进小麦根系伸长,高浓度则抑制;与黄体酮对小麦根系伸长生长的影响趋势相似。

图3

图3   外源葡萄糖对小麦根系生长的影响

Fig.3   Effects of exogenous glucose on wheat root elongation


2.4 外源黄体酮和葡萄糖交互作用对小麦根长的影响

黄体酮对小麦根长的调控与葡萄糖代谢有关(图4)。与PG0处理相比,10 μmol/L葡萄糖(PG1和PG2)处理促进小麦幼苗根系伸长,10 000 μmol/L葡萄糖(PG3和PG4)处理抑制根系伸长。PG2处理在10 μmol/L葡萄糖(促进根系伸长)处理的同时,加入0.1 μmol/L黄体酮(抑制根系伸长)可显著抑制外源葡萄糖对根系伸长的促进作用,PG2较PG1处理的根长降低了14.5%。相反,PG4处理在10 000 μmol/L葡萄糖(抑制根系伸长)处理同时,加入0.001 μmol/L黄体酮(促进根系伸长)可缓解外源葡萄糖对根系伸长的抑制作用,PG4处理的根长较PG3处理增加了40.7%。结合黄体酮可以显著影响根系中葡萄糖含量及PFK活性的试验结果(图2)分析得出,黄体酮对小麦根系伸长的调控与葡萄糖代谢途径密切相关。

图4

图4   外源黄体酮和葡萄糖互作处理对小麦根系生长的影响

“*”表示处理间在P < 0.05水平差异显著,下同。

Fig.4   Effects of exogenous progesterone and glucose on root elongation of wheat

“*”indicates significant difference among treatments at P < 0.05 level, the same below.


2.5 外源黄体酮和葡萄糖交互作用对小麦根系中葡萄糖含量及PFK活性的影响

图5所示,PG1处理下幼苗根系中的葡萄糖含量降低,PFK活性增加;PG3和PG4处理下幼苗根系中的葡萄糖含量升高,PFK活性受到抑制。PG1和PG3处理下,小麦幼苗根系中葡萄糖含量和PFK活性变化趋势与单独黄体酮处理下(图2)的趋势基本一致。

图5

图5   外源黄体酮和葡萄糖互作处理对小麦根系中葡萄糖含量和PFK活性的影响

Fig.5   Effects of exogenous progesterone and glucose on glucose content and PFK activity in wheat roots


PG2处理下,加入0.1 μmol/L黄体酮可显著促进10 μmol/L葡萄糖处理下小麦根系中葡萄糖的积累,约较PG0处理提高了78.4%;PG1和PG2处理间葡萄糖含量差异显著。PG4处理中,0.001 μmol/L黄体酮的加入显著抑制了10 000 μmol/L葡萄糖处理下根系葡萄糖的积累,降幅为50.6%。另外,PG2处理中0.1 μmol/L黄体酮的加入显著抑制了10 μmol/L葡萄糖处理下小麦根系PFK活性的增加,降至PG1处理下的60.5%。而在PG4处理下,0.001 μmol/L黄体酮的加入有效解除了外源葡萄糖对PFK活性的抑制,PFK活性显著提高到PG3处理下的1.66倍。

3 讨论

苗期是小麦生长发育的关键起始阶段,也是根系形态建成和生理功能完善的核心时期,根系生长状况直接决定麦苗长势、群体结构和产量[19]。在根系诸多性状中,根伸长决定根系入土深度、空间分布范围与养分水分吸收能力,是小麦抵御干旱、低温和盐碱等非生物逆境胁迫的重要基础[20]。大量研究[21-22]表明,苗期根系伸长充分且下扎较深的小麦植株,其水肥捕获能力强、分蘖成穗率更高以及抗倒伏性更优,对实现稳产高产具有显著促进作用。

植物根系的生长发育受多种植物激素调控,如生长素、细胞分裂素、赤霉素、脱落酸、乙烯和油菜素甾醇等[23],这些激素相互作用,共同调控根系生长[24]。黄体酮被认为是植物体内的一种候选激素,探讨其对植物根系的调控作用具有重要意义。本研究表明,外源黄体酮对小麦根系伸长生长的影响表现为低浓度促进、高浓度抑制,这与前人在拟南芥[17]和向日葵[1]中的研究结果相似。前人[25-27]研究表明,生长素与葡萄糖对植物根系的调控存在交互作用,高浓度葡萄糖通过降低根分生组织中的生长素水平,最终抑制拟南芥根系生长[28-29];而适宜浓度的葡萄糖与生长素共同处理,可显著增强单独处理对苹果根长的促进作用[30]。本研究在上述试验结果的基础上,进一步细化了葡萄糖处理浓度,发现低浓度葡萄糖促进小麦根系伸长,在10 μmol/L葡萄糖浓度处理下,小麦根系平均长度达到最大值,随后随葡萄糖浓度升高,根系伸长受到抑制。此外,本研究发现黄体酮处理对小麦根系伸长的影响趋势与葡萄糖处理高度吻合,且黄体酮可显著改变小麦根系中的葡萄糖含量。低浓度黄体酮处理降低根系葡萄糖含量,而高浓度处理则显著增加其含量。由此推测,低浓度黄体酮促进葡萄糖代谢,而高浓度黄体酮则抑制葡萄糖代谢,黄体酮可能通过调控根系葡萄糖含量影响小麦根系的伸长生长。

为明确黄体酮通过调控根系葡萄糖含量影响小麦根系伸长生长的潜在机制,本试验开展了进一步研究。葡萄糖的氧化代谢始于糖酵解途径,PFK是该途径的关键限速酶,其活性直接决定糖酵解速率。本研究检测了黄体酮处理下小麦根系中PFK的活性变化,结果表明低浓度黄体酮处理可显著提升小麦根系中PFK的活性,而高浓度黄体酮则显著抑制其活性。黄体酮与葡萄糖的交互试验表明,葡萄糖处理下根系PFK活性的变化趋势与黄体酮处理一致,低浓度葡萄糖显著增强PFK活性,高浓度则显著抑制。同时,高浓度黄体酮可抑制低浓度葡萄糖对PFK的诱导作用,而低浓度黄体酮可缓解高浓度葡萄糖对PFK的抑制效应。基于上述结果,推测低浓度黄体酮通过增加PFK活性和加速糖酵解进程,从而降低根系葡萄糖含量;而高浓度黄体酮则抑制PFK活性并减缓糖酵解进程,导致根系葡萄糖积累、含量升高。本研究明确了外源黄体酮对小麦根系伸长生长的影响,并初步分析了其调控机制,为进一步探明黄体酮调控小麦根系伸长的调控路径提供了理论参考。

4 结论

外源黄体酮可显著影响小麦根系的伸长生长,表现为低浓度促进、高浓度抑制。0.001 μmol/L和0.01 μmol/L黄体酮处理下,小麦根长较对照分别增加了15.3%和9.0%。大田生产中应用低浓度黄体酮进行浸种或根施可有效促进小麦根系伸长,提高抗逆性。黄体酮可能通过调控PFK活性影响糖酵解进程,进而影响小麦根系中的葡萄糖含量,实现对根系伸长生长的调控。低浓度黄体酮通过提升PFK活性加速糖酵解进程,促进葡萄糖代谢,降低根系葡萄糖含量,最终促进根系伸长;高浓度黄体酮抑制PFK活性,减缓糖酵解进程,导致根系葡萄糖积累,抑制根系伸长。

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