新疆农业科学, 2023, 60(3): 624-632 DOI: 10.6048/j.issn.1001-4330.2023.03.013

园艺特产·生理生化

辣椒核雄性不育系pby-1形态学及生理生化特性分析

裴红霞,1, 李生梅2, 武旭霞3, 耿世伟2, 赖黎丽4, 高晶霞,1, 董心久,2

1.宁夏农林科学院园艺研究所,银川 750002

2.新疆农业科学院经济作物研究所,乌鲁木齐 830091

3.呼和浩特职业学院,呼和浩特 010070

4.新疆天椒红安农业科技有限责任公司,新疆石河子 832011

Morphological Observation and Physio-Biochemical Characteristics of Hot Pepper Genic Male - Sterile Line pby-1

PEI Hongxia,1, LI Shengmei2, WU Xuxia3, GENG Shiwei2, LAI Lili4, GAO Jingxia,1, DONG Xinjiu,2

1. Institute of Horticulture, Ningxia Academy of Agriculture and Forestry Science, Yinchuan 750002, China

2. Research Institute of Economic Crops, Xinjiang Academy of Agricultural Sciences, Urumqi 830091, China

3. Hohhot vocational college, Huhehaote, 010070, China

4. Xinjiang Tianjiao Hong'an Agricultural Technology Co., Shihezi Xinjiang 832011, China

通讯作者: 高晶霞(1984-), 女, 宁夏人,助理研究员, 研究方向为作物遗传育种, (E-mail)774350762@qq.com;董心久(1980-), 男, 山东人, 副研究员, 研究方向为作物栽培与生理, (E-mail)80416913@qq.com

收稿日期: 2022-07-30  

基金资助: 宁夏自然科学联合基金(2021AAC03264)
宁夏回族自治区农业科技自主创新资金(NGSB-2021-8-01)
新疆维吾尔自治区优秀青年科技人才培养项目(2018Q16)

Corresponding authors: GAO Jingxia (1984-), female, from ningxia, research assistant, vegetable genetics and breeding research, (E-mail)774350762@qq.com;DONG Xinjiu (1980-), male, from Shandong, research asociate, crop cultivation and physiological research, (E-mail)80416913@qq.com

Received: 2022-07-30  

Fund supported: Ningxia Natural Science Union Foundation(2021AAC03264)
Agricultural Science and Technology Independent Innovation Fund of Ningxia Hui Autonomous Region(NGSB-2021-8-01)
Outstanding Young Scientific and Technological Talents Training Program of Xinjiang Uygur Autonomous Region(2018Q16)

作者简介 About authors

裴红霞(1980-),女,山西人,助理研究员,研究方向为作物遗传育种,(E-mail)810444147@qq.com

摘要

【目的】分析辣椒(Capsicum annuum L.)雄性不育系的形态学及生理生化的特征,为研究羊角椒雄性不育的遗传机制提供理论参考。【方法】以羊角椒核雄性不育系pby-1和保持系PBY-1为材料,观察花蕾7个时期的发育情况及盛花期的花器官,测定盛花期花粉的生活力及减数分裂、四分体及单核3个时期花蕾的超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)和丙二醛(MDA)的含量,分析保护酶活性的变化,比较赤霉素(GA3)、生长素(IAA)、玉米素核苷(ZR)、茉莉酸(JA)及脱落酸(ABA)含量差异及不同内源激素的平衡关系。【结果】与野生型PBY-1相比,不育系只有花器官结构与其不同,花药干瘪,花粉无生活力。在羊角椒花蕾发育的3个时期,不育系与可育系之间在保护酶活性与内源激素含量上存在差异,其中,不育系SOD活性始终显著低于保持系,而CAT活性始终显著高于保持系,POD活性在减数分裂期和四分体时期显著低于保持系,MDA含量只有在单核时期显著高于保持系;四分体及单核时期GA3含量、IAA含量高于保持系,ZR含量始终高于保持系,减数分裂期JA含量明显低于保持系,四分体时期IAA含量明显高于保持系;不育系和保持系IAA/ABA、IAA/GA3、IAA/JA、IAA/ZR、ABA/GA3、ABA/ZR的变化趋势不一致,且IAA/ABA的比值高于其他内源激素含量的比值。【结论】不育系保护酶活性的变化和内源激素含量的异常以及激素比值的变化可能影响了减数分裂、四分体及单核3个时期的花药内源物质和花粉活力,导致羊角椒不育系花药中花粉发育受阻、形成败育。

关键词： 辣椒; 雄性不育; 形态学; 抗氧化物; 内源激素

Abstract

【Objective】 This project aims to explore the morphological, physio-biochemical characteristics of male sterile lines of pepper (Capsicum annuum L.) in the hope of providing a theoretical reference for revealing the genetic mechanism of male sterility of hot pepper.【Methods】 The pepper nuclear male sterility line pby-1 and the maintainer line PBY-1 were used as test materials.The development of flower buds at 7 stages and the flower organs in the full blooming period were observed, and the viability of the pollen in the full blooming period was measured.The contents of superoxide dismutase (SOD), peroxidase (POD), catalase (CAT) and malondialdehyde (MDA) in flower buds at the three stages of meiosis number, tetrad and mononuclear were analyzed.The contents of gibberelli cacid (GA3), acetic acid (IAA), zeatin riboside (ZR), jasmonic acid (JA) and abscisic acid (ABA) and the balance relationship between different endogenous hormones were compared.【Results】 Compared with the wild-type PBY-1, the sterile line only differed from its floral organ structure, the anthers were shriveled, and the pollen had no viability.In the three stages of flower bud development of hot pepper, there were differences in protective enzyme activity and endogenous hormone content between the sterile line and the maintainer line.Among them, the SOD activity of the sterile line was always significantly lower than that of the maintainer line, while the CAT activity was always significantly higher than that of the maintainer line, POD activity was significantly lower than that of the maintainer line during the meiotic and tetrad period, and the MDA content was significantly higher than that of the maintainer line only in the mononuclear period.In the three periods, there were differences in protective enzyme activity and endogenous hormone content between the sterile line and the maintainer line in the development of hot pepper.Among them, the SOD activity of the sterile line was always significantly lower than that of the maintainer line, while the CAT activity was significantly higher than that of the maintainer line.The POD activity was significantly lower than that of the maintainer line during the meiotic and tetrad period, and the MDA content was only significantly higher than maintainer in the mononuclear period.Content GA3 and IAA in tetrad and mononuclear period in sterile line were higher than that of the maintainer, ZR content was always higher than that of the maintainer, JA content in meiosis stage was significantly lower than that of the maintainer, IAA content in tetrad period was significantly higher in the maintainer line.The change trends of IAA/ABA, IAA/GA3, IAA/JA, IAA/ZR, ABA/GA3, ABA/ZR between sterile line and the maintainer line were inconsistent.The ratio content of IAA/ABA was higher than those of other endogenous hormones.【Conclusion】 Therefore, it is speculated that the changes of the protective enzyme activity, the abnormalcontent of endogenous hormone and the change in the hormone ratio in sterile line may affect anther endogenous substances and pollen vitality in the three stages of meiosis, tetrad, and mononucleus, resulting in the development of pollen in the anthers of the male sterile line in croissant pepper is hindered and the cause of abortion is formed.

Keywords： Capsicum annuum L.; male sterility; morphology; antioxidant; endogenous hormone

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本文引用格式

裴红霞, 李生梅, 武旭霞, 耿世伟, 赖黎丽, 高晶霞, 董心久. 辣椒核雄性不育系pby-1形态学及生理生化特性分析[J]. 新疆农业科学, 2023, 60(3): 624-632 DOI:10.6048/j.issn.1001-4330.2023.03.013

PEI Hongxia, LI Shengmei, WU Xuxia, GENG Shiwei, LAI Lili, GAO Jingxia, DONG Xinjiu. Morphological Observation and Physio-Biochemical Characteristics of Hot Pepper Genic Male - Sterile Line pby-1[J]. Xinjiang Agricultural Sciences, 2023, 60(3): 624-632 DOI:10.6048/j.issn.1001-4330.2023.03.013

0 引言

【研究意义】辣椒(Capsicum annuum L.)属于茄科[1],与常规品种相比,辣椒杂种优势的增产作用体现的非常明显[2]。雄性不育的利用是杂种优势利用的重要途径之一,目前,辣椒杂交种主要依靠人工去雄授粉,制种成本高[3]。雄性不育是杂种优势利用的重要途径之一,省时省力,能提高种子纯度[4]。研究辣椒雄性不育遗传机制,对解析辣椒雄性不育形成的机理有重要意义。【前人研究进展】已有对植物雄性不育的细胞形态学、生理生化特性和分子生物学等多方面领域进行的研究报道[5]。其中,细胞生物学是研究植物雄性不育的基础[6]。植物雄性不育的生理生化特性与表型变异也有所联系,生长发育的成分包括酶、营养物质和生长物质等,多方面互相协同才能正常生长[7]。花粉的发育与雄蕊尤其是花药密切相关,若发生物质代谢紊乱、能量亏缺或者激素和酶的异常调控都可能会影响雄性不育[8,9]。植物在胁迫或核质遗传不协调等引起雄性不育的情况下,可能会产生活性氧,进而对细胞膜系统形成伤害,而抗氧化酶系统(SOD、POD和CAT等)能有效清除活性氧自由基,使植物体内活性氧的产生与清除达到动态平衡的关系[10,11],随着小孢子的发育,甘蓝型油菜核雄性不育系花蕾,SOD、POD和CAT活性显著低于保持系[12]。内源激素是雄性不育发生的关键调节的重要因子[13],花椰菜不育系花蕾中的GA和ABA含量总体呈上升趋势且显著高于保持系[14]。【本研究切入点】近来年对甜瓜(Cucumis melo)[15]、陆地棉(Gossypium hirsutum L.)[16]、厚轴茶(Camellia crassicolumna)[17]、西瓜(Citrullus lanatus var.lanatus)[18]等多种植物开展了雄性不育生理生化特征的研究,分析核雄性不育与抗氧化酶和内源激素的关系,发现不育系和保持系相关生理生化指标的含量存在较大差异。而小孢子的发育与花蕾外部形态密切相关,可能花蕾的大小直接决定了小孢子的发育时期[19]。植物体内含有较高的MDA也是造成败育的重要原因,如甜瓜雄性不育系的雄蕊发育过程中各时期不育系MDA含量高于保持系。目前对辣椒雄性不育酶活性的研究相对较少。需分析辣椒(Capsicum annuum L.)雄性不育系的形态学及生理生化的特征。【拟解决的关键问题】以细胞核雄性不育系突变体pby-1,保持系PBY-1为材料,研究雄性败育发生的时期和特点,测定花蕾发育关键时期的生理特性,分析内源激素比值及pby-1细胞核雄性不育系败育的原因,为辣椒杂种优势的利用提供理论依据。

1 材料与方法

1.1 材料

供试材料为细胞核雄性不育系pby-1,保持系PBY-1,材料由宁夏农林科学院园艺研究所辣椒课题组提供。

田间试验于2020年3月25日育苗,5月7日单株定植于宁夏园林场试验基地(E 106.060°,N 38.620°),露地栽培。

1.2 方法

1.2.1 试验设计

随机区组排列(30 m2/小区),设3次重复,株距为45 cm,行距为65 cm,常规方法进行田间管理。调查取样全部在宁夏农林科学院园林场基地内进行。

1.2.2 测定指标
1.2.2.1 花器官观察以及花粉生活力

在2020年盛花期,于晴天09∶00~10∶00取不育株pby-1和可育株PBY-1不同时期的花蕾,根据花蕾形态特征与花粉发育时期将花蕾分为7级[20],分别取造孢时期(Ⅰ)、花粉母细胞时期(Ⅱ)、减数分裂时期(Ⅲ)、四分体时期(Ⅳ)、单核小孢子时期(Ⅴ)、双核小孢子时期(Ⅵ)和成熟花粉粒时期(Ⅶ)的花蕾,采用游标卡尺测定各时期花蕾纵横径,拍照,花粉活力测定采用黄炜花粉离体萌发测定法[21]。

1.2.2.2 生理生化指标测定

取减数分裂时期(Ⅲ)、四分体时期(Ⅳ)及单核时期(Ⅴ)的花蕾测定各项生理指标,实验设置3个重复。利用SpectraMaxi3x酶标仪测定丙二醛(MDA)、超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT);利用LC210高效液相色谱仪测定了赤霉素(GA3)、生长素(IAA)、玉米素核苷(ZR)、茉莉酸(JA)和脱落酸(ABA),具体参照实验指导书[22]。

2 结果与分析

2.1 羊角椒不同时期花蕾形态特征及动态发育

研究表明,不育系pby-1与保持系PBY-1的花蕾进程基本一致,分为7个阶段,除了不育系pby-1的花蕾在成熟花粉粒时期(S7)出现干瘪外,其余各时期不育系与保持系之间无明显差异。羊角椒在造孢时期(Ⅰ)、花粉母细胞时期(Ⅱ)、减数分裂时期(Ⅲ)、四分体时期(Ⅳ)、单核小孢子时期(Ⅴ)、双核小孢子时期(Ⅵ)和成熟花粉粒时期(Ⅶ)花蕾纵横径大小,基本无明显差异。 图1,表1

图1

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图1   羊角椒不同时期花蕾的发育过程

注:S1~S7:不育系Ⅰ-Ⅶ时期;F1~F7:保持系Ⅰ-Ⅶ时期

Fig.1   The process in during bud development of hot pepper

Note:S1-S7:sterile line Ⅰ-Ⅶ period;F1-F7:maintainer line Ⅰ-Ⅶ period


表1   羊角椒不同时期花蕾的形态特征与大小

Tab.1  The morphological characteristics and size in during bud development of hot pepper

材料
Materials		小孢子
发育时期
Microspore
development
time		花蕾形态特征
Morphological characteristics flower buds		花蕾纵横径
Veritical and horizontal
diameter of flower bud
纵径
Longitudinal
diameter(mm)		横径
Transverse
diameter(mm)
pby-1S1花蕾小,花萼紧裹花冠,深绿色。2.49~2.982.07~2.63
S2花蕾比S1期花蕾微大,花萼裹着花冠,深绿色。2.99~4.492.64~3.59
S3花萼上端微张开,深绿色,花冠绿色,花冠与花萼齐平。4.50~5.873.60~4.10
S4花萼上端张开,深绿色,花冠绿色,花冠约占花萼长的1/4。5.88~6.524.11~4.86
S5花冠绿色,伸出花萼部分的长度短于花萼,约占花萼长的1/2。6.53~7.234.87~5.54
S6花冠浅绿色,伸出花萼部分的长度约与花萼等长。7.22~8.895.55~5.90
S7花冠白绿色,大小充分膨大。8.90~11.985.91~6.40
PBY-1F1花蕾小,花萼紧裹花冠,深绿色。2.57~2.932.15~2.84
F2花蕾比F1期花蕾微大,花萼裹着花冠,深绿色。2.94~4.812.85~3.73
F3花萼上端微张开,深绿色,花冠绿色,花冠与花萼齐平。4.82~5.753.72~4.08
F4花萼上端张开,深绿色,花冠绿色,花冠约占花萼长的1/4。5.76~6.664.09~4.94
F5花冠绿色,伸出花萼部分的长度短于花萼,约占花萼长的1/2。6.67~7.284.95~5.76
F6花冠浅绿色,伸出花萼部分的长度约与花萼等长。7.29~8.085.77~5.91
F7花冠白绿色,大小充分膨大。8.09~11.845.92~6.80

注:S1~S7:不育系Ⅰ-Ⅶ时期;F1~F7:保持系Ⅰ-Ⅶ时期

Note:S1-S7:sterile line Ⅰ-Ⅶ period;F1-F7:maintainer line Ⅰ-Ⅶ period

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2.2 羊角椒花器形态特征及花粉生活力测定

研究表明,野生型PBY-1植株的花粉粒饱满,平均花粉萌发率为81.14%,花粉管的伸长能力很强,花粉育性也正常,而突变体pby-1植株花粉并没有生活力。不育系pby-1花蕾发育、开花习性等表现正常,雌蕊不仅具有正常发育的能力还具有异交授粉受精的能力,与野生型PBY-1相比,唯独花器结构不同,花器官退化较轻,花冠外表正常,花药稍瘪,花药中的花粉量极少甚至无花粉,授粉时不用去掉花药就可以杂交授粉,突变体pby-1植株盛花期全株落花现象严重,会有部分单性结实。图2,图3

图2

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图2   PBY-1和pby-1的花粉萌发比较

Fig.2   Comparison of pollen germination between PBY-1 and pby-1


图3

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图3   A PBY-1和pby-1的花器形态比较B PBY-1和pby-1的果实形态比较

Fig.3   A: Comparison of floral morphology of PBY-1 and pby-1B Comparison of fruit morphology of PBY-1 and pby-1


2.3 羊角椒不同时期花蕾保护酶活性的变化

研究表明,在花蕾发育过程中,pby-1、PBY-1花蕾的SOD活性均呈先下降后上升趋势,在四分体时期出现最低值,分别为307.28和333.55 U/g,且pby-1花蕾的SOD活性从减数分裂时期至单核期均显著低于相应的PBY-1花蕾(P<0.05);pby-1花蕾的POD活性总体呈现上升趋势,而PBY-1花蕾的POD活性则是呈现先上升后下降的趋势,在减数分裂期、四分体时期PBY-1花蕾的POD活性均显著高于相应的pby-1(P<0.05);从减数分裂期至单核期,PBY-1的花蕾CAT活性先降低后升高,变化比较平缓,无明显差异,而pby-1的花蕾CAT活性变化趋势与PBY-1不同,呈现出不断下降趋势,且从减数分裂时期至单核时期pby-1的花蕾CAT活性始终显著低于PBY-1(P<0.05);花药发育过程中,pby-1、PBY-1中MDA含量都逐步降低,在减数分裂时期,pby-1、PBY-1中MDA含量几乎无差异,在四分体时期,pby-1花蕾MDA含量超过了PBY-1,在单核期,pby-1花蕾MDA含量显著高于PBY-1(P<0.05)。图4

图4

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图4   羊角椒不同时期花蕾保护酶活性变化

注:Ⅲ、Ⅳ和Ⅴ分别表示减数分裂时期、四分体时期及单核时期的花蕾;不同小写字母表示差异在P<0.05水平达到显著水平。下同

Fig.4   Changes in the activity of protective enzymes in during bud development of hot pepper

Note:Ⅲ, Ⅳ, and Ⅴ indicate the flower buds in meiosis number, tetrad and mononuclear respectively; different letters shows significantly different at P<0.05 levels,the same as below


2.4 辣椒不同时期花蕾内源激素含量的变化

研究表明,在花蕾从减数分裂时期至单核期发育的整个过程中,不育系pby-1花蕾的GA3含量先升高后降低,而PBY-1花蕾的GA3含量逐渐降低,pby-1花蕾的GA3含量在减数分裂期、四分体期及单核期分别是PBY-1的47.7%、105.8%、166.9%。pby-1与PBY-1花蕾的IAA含量变化趋势相同,均呈现出逐渐降低的趋势,在减数分裂时期,pby-1花蕾的IAA含量与PBY-1相当,而到了四分体期、单核期pby-1花蕾的IAA含量明显高于PBY-1,分别是PBY-1花蕾IAA含量的121.1%、183.4%。pby-1与PBY-1花蕾的ZR含量变化趋势相同,均呈现出先升高后降低的趋势,且在3个时期中,不育系pby-1花蕾的ZR含量均高于PBY-1,不育系pby-1花蕾的ZR含量分别占PBY-1的146.2%、123.1%及129.3%。不育系pby-1花蕾的JA含量总体呈现上升趋势,而PBY-1的JA含量总体呈现下降趋势,在减数分裂期,PBY-1花蕾的JA含量明显高于pby-1,是其花蕾JA含量的185.7%。不育系pby-1与PBY-1花蕾的ABA含量变化趋势明显不同,pby-1花蕾的ABA含量总体呈先下降后上升的趋势,而PBY-1花蕾的ABA含量总体呈先上升后下降趋势,在四分体时期,PBY-1花蕾的ABA含量明显高于pby-1,是其花蕾ABA含量的215.4%。图5

图5

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图5   羊角椒不同时期花蕾内源激素含量的变化

Fig.5   Changes of endogenous hormones contents in during bud development of hot pepper


2.5 辣椒不同时期花蕾内源激素之间的平衡关系

研究表明,IAA/ABA的比值高于其他内源激素含量的比值,且不育株pby-1只有在减数分裂时期的比值低于可育株;对于IAA/GA3,3个时期不育株pby-1的比值均高于可育株的比值,四分体时期和单核期的比值相差不大,不育株与可育株分别差0.15和0.16;对于IAA/JA,发现不育株的比值较高,3个时期不育株pby-1的比值呈下降趋势,而可育株的比值是先上升后下降;对于IAA/ZR,在减数分裂时期不育株与可育株的比值均最高,其他2个时期的比值都有所下降;对于ABA/GA3,不育株pby-1的比值先下降后上升,而可育株的比值先上升后下降,2个材料比值变化趋势相反;对于ABA/ZR,不育株pby-1在减数分裂时期比值最高,四分体时期与单核期激素的比值变化平缓,可育株的比值最高是在四分体时期,其余2个时期的变化趋势与不育株走势相反。表2

表2   羊角椒不同时期花蕾内源激素含量比值变化

Tab.2  Changes in the ratio of endogenous hormone content in during bud development of hot pepper

材料
Material		发育时期
Bud stage		IAA/ABAIAA/GA3IAA/JAIAA/ZRABA/GA3ABA/ZR
pby-1减数分裂2.202.460.540.891.120.41
四分体时期3.951.290.450.680.330.17
单核时期3.361.440.370.641.430.19
PBY-1减数分裂4.111.280.291.300.310.32
四分体时期1.511.110.350.690.740.45
单核时期2.841.270.210.450.450.16

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3 讨论

细胞核不育的根本原因是花药发育异常,从而产生无活性的花粉粒[23]。目前在拟南芥(Arabidopsis thaliana)[24]、小麦(Triticum aestivum L.)[25]、油菜(Brassica campestris L.)[26]、水稻(Oryza sativa L.)[27]等植物均发现雄性不育的原因是花药内壁绒毡层发育异常引起。对羊角椒进行调查发现,不育系pby-1和保持系PBY-1的发育进程基本一致,而在发育后期,不育系花蕾与花药均呈现稍瘪状态,且无花粉,羊角椒花药的发育及花蕾的外部形态与其雄性不育性密切相关。

SOD、POD和CAT等保护酶活性与植物雄性不育现象密切相关,如棉花细胞质雄性不育系花蕾发育过程中CAT活性在败育前低于保持系,POD活性在败育前高于保持系[16]。厚轴茶不育系花蕾发育过程中不育系POD活性始终显著高于保持系,不育系在单核期和花粉成熟期SOD和CAT活性都显著低于保持系[17]。辣椒品种不同雄性不育系材料与保持系之间3种酶活性变化也存在一定的差异[28]。试验研究与前人部分结果一致,羊角椒花蕾发育3个时期中,不育系SOD活性显著低于保持系,不育系POD活性只有在单核时期略高于保持系,不育系CAT活性显著高于保持系,不育系MDA含量高于保持系且在单核时期显著高于保持系。酶类活性的变化致使花粉粒内新陈代谢紊乱从而造成雄性不育。

内源激素间的平衡与植物雄性不育相关密切,IAA含量的增多,乙烯的过度产生,ABA含量的增加,以及GA3和ZR含量的降低,内源JA含量减少都可能导致植物雄性不育[29]。如水稻不育系在四分体时期和花粉成熟期GA的含量低于保持系[30],而花椰菜不育系在这2个时期GA的含量高于保持系[14],不同不育系材料GA含量的变化并不一致。百合花药在各个发育阶段的IAA含量均表现为高于保持系,在四分体时期不育系ZR和GA4含量低于保持系,而JA和ABA含量高于保持系[8]。芝麻在花蕾发育阶段中,IAA含量始终显著低于保持系,JA和ABA含量始终高于保持系[31]。研究羊角椒花蕾发育3个时期中,减数分裂时期不育系GA3含量低于保持系,而ZR、JA和ABA含量高于保持系;四分体时期不育系GA3、IAA和ZR含量高于保持系,JA和ABA含量低于保持系;在单核期不育系GA3、IAA、ZR、JA和ABA含量高于保持系。减数分裂时期GA3含量的下降,四分体时期JA含量的减少,ABA含量的增加,导致羊角椒雄性不育。

植物雄性不育除了受保护酶活性和内源激素含量影响之外,也可能受激素间相互协作或者相互拮抗共同起作用的结果,如大白菜不育系花蕾发育过程中ABA/GA3和IAA/GA3比值都明显大于保持系,不育系与保持系IAA/ABA比值呈先下降后上升趋势,不育系与保持系IAA/ZR比值变化趋势则不同[32]。辣椒胞质雄性不育系花蕾发育过程中IAA/ZR、ABA/ZR以及IAA/GA3的比值均高于保持系[33]。研究得出的ABA/GA3、IAA/GA3的变化趋势与前人研究结果一致,而IAA/ABA、IAA/JA、IAA/ZR和ABA/ZR的变化规律略有不同,不同雄性不育的材料内源激素的比值变化趋势不同,不同材料败育的机理存在差异。

4 结论

羊角椒小孢子从减数分裂期至双核期的发育过程中,不育系花蕾中CAT活性、MDA含量、ZR含量、IAA/GA3和IAA/JA一直高于可育系,尤其在双核期,不育系花蕾中CAT活性和MDA含量分别达到了663.77 nmol/(min·g)、19.30 nmol/g,是可育系花蕾的165.55%、113.13%,CAT无法及时清除花药细胞中产生的ROS,而高水平的ROS会破坏脂膜系统,造成细胞代谢紊乱,引起败育。在羊角椒花蕾发育的过程中,保护酶活性的变化和内源激素含量的异常以及激素间相互协作或者相互拮抗可能影响了花药内源物质和花粉活力,导致羊角椒不育系花药中花粉发育受阻、败育。

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为了探究内源激素在花椰菜温敏雄性不育系GS-19育性调控中的作用,以花椰菜温敏雄性不育系GS-19为材料,采用高效液相色谱法,对花椰菜温敏雄性不育系的不育株和可育株花蕾不同发育时期及叶片中赤霉素(GA)、生长素(IAA)、脱落酸(ABA)及玉米素核苷(ZR)的动态变化进行比较分析。结果表明,花椰菜温敏雄性不育系花蕾和叶片发育过程中,不育株和可育株内源激素GA、IAA、ABA和 ZR的变化均存在明显差异。不育株花蕾中的GA和ABA含量总体呈上升趋势,GA含量在造孢时期、四分体时期及花粉成熟期的含量均显著高于可育株,分别为281.54、355.37和350.13 &#x003bc;g&#x000b7;g<sup>-1</sup>,比可育株花蕾高45.1%、210.4%和54.5%,而ABA含量只在四分体时期和花粉成熟期显著高于可育株,分别为2.22 &#x003bc;g&#x000b7;g<sup>-1</sup>和2.88 &#x003bc;g&#x000b7;g<sup>-1</sup>,比可育株花蕾高82%和35.2%;不育株花蕾中的IAA含量先降后升,在造孢时期和花粉成熟期显著高于可育株,分别为1.97 &#x003bc;g&#x000b7;g<sup>-1</sup>和2.55 &#x003bc;g&#x000b7;g<sup>-1</sup>,比可育株花蕾高52.7%和82.1%,但不育株花蕾的ZR含量先升后降,在四分体和花粉成熟期显著高于可育株,分别为320 &#x003bc;g&#x000b7;g<sup>-1</sup>和170 &#x003bc;g&#x000b7;g<sup>-1</sup>,比可育株花蕾高580.9%和243.9%。不育株的IAA/ABA呈 &#x0201c;V&#x0201d;字型变化,在四分体时期比值最低,而GA/ABA和ZR/ABA呈倒 &#x0201c;V&#x0201d;字型的变化,在四分体时期比值最高。叶片中ABA和ZR未检出,GA和IAA含量不育株显著高于可育株。说明内源激素含量升高和多种激素间的平衡关系被打破,比例失调,阻碍了小孢子的正常发育,导致花椰菜温敏雄性不育系GS-19花粉败育。本研究结果对揭示内源激素在雄性败育过程中可能存在的作用机制具有重要意义,为进一步揭示花椰菜温敏雄性不育的遗传机制积累了资料。

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DOI      [本文引用: 1]

以西瓜雄性不育系&lsquo;Se18&rsquo;为试材，研究开花期间花蕾和叶片中抗氧化酶活性和内源激素含量的变化。结果表明：不育株雄花花蕾在整个发育时期，超氧化物歧化酶（SOD）和过氧化物酶（POD）活性升高，过氧化氢酶（CAT）活性降低；可育株中CAT 活性升高，SOD 和POD 活性降低。不育株和可育株雄花花蕾中的生长素（IAA）、脱落酸（ABA）、赤霉素（GA3）、玉米素核苷（ZR）、茉莉酸（JA）和油菜素内酯（BR）及异戊烯基腺嘌呤核苷（IPA）等7 种内源激素含量变化趋势不同，且含量差异明显。其中，不育株中IAA、GA3、BR 和IPA 含量降低，JA 含量升高；可育株中JA、BR、GA3 和IPA 含量降低，IAA 和ZR 含量升高。不育株叶片中的IAA、ZR 和BR 含量显著下降，ABA 和JA 含量显著上升；IAA/ABA、IAA/GA3、IAA/JA、IAA/ZR、IAA/BR、IAA/IPA 及ABA/GA3、ABA/ZR 在不育株和可育株间的变化趋势不一致，且差异较大。因此推测上述抗氧化酶活性变化和内源激素含量异常可能与西瓜雄性不育的发生密切相关。

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DOI      [本文引用: 1]

Pollen and microspore development are essential steps in the life cycle of all land plants that generate male gametes. Within flowering plants, pollen development occurs inside of the anther. Here, we report the identification of two class III peroxidase-encoding genes, PEROXIDASE9 (PRX9) and PRX40, that are genetically redundant and essential for proper anther and pollen development in Arabidopsis (Arabidopsis thaliana). Arabidopsis double mutants devoid of functional PRX9 and PRX40 are male sterile. The mutant anthers display swollen, hypertrophic tapetal cells and pollen grains, suggesting disrupted cell wall integrity. These phenotypes lead to nearly 100%-penetrant pollen degeneration upon anther maturation. Using immunochemical and biochemical approaches, we show that PRX9 and PRX40 likely cross-link extensins to contribute to tapetal cell wall integrity during anther development. This work suggests that PRX9 and PRX40 encode Arabidopsis extensin peroxidases and highlights the importance of extensin cross-linking during pollen development.

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[本文引用: 1]

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