1 材料和方法

1.1 材料

1.2 材料种植

1.3 田间育性鉴定

1.4 室内花粉镜检

1.5 统计分析

2 结果与分析

2.1 恢保关系测定

2.2 育性恢复组合F₂群体的育性表现

图1

图2

2.3 双交群体的育性表现

图3

图4

3 讨 论

3.1 同质异核不育系对相同测验系的育性恢复反应不同,暗示不育系核背景可能参与调控育性恢复表现

3.2 微效基因通过累加达到某一个阈值也可以实现对玉米CMS-C不育系的育性恢复

3.3 环境不仅对玉米CMS-C弱恢复系的恢复作用产生影响,有时对强恢复系的恢复作用也会产生影响

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。

参考文献 View Option 原文顺序
文献年度倒序
文中引用次数倒序
被引期刊影响因子

[1] Chen LT, Liu YG . Male sterility and fertility restoration in crops
Annu Rev Plant Biol, 2014,65(1):579-606.
URLPMID:24313845 [本文引用: 1]
Abstract In plants, male sterility can be caused either by mitochondrial genes with coupled nuclear genes or by nuclear genes alone; the resulting conditions are known as cytoplasmic male sterility (CMS) and genic male sterility (GMS), respectively. CMS and GMS facilitate hybrid seed production for many crops and thus allow breeders to harness yield gains associated with hybrid vigor (heterosis). In CMS, layers of interaction between mitochondrial and nuclear genes control its male specificity, occurrence, and restoration of fertility. Environment-sensitive GMS (EGMS) mutants may involve epigenetic control by noncoding RNAs and can revert to fertility under different growth conditions, making them useful breeding materials in the hybrid seed industry. Here, we review recent research on CMS and EGMS systems in crops, summarize general models of male sterility and fertility restoration, and discuss the evolutionary significance of these reproductive systems.

[2] Beckett JB . Classification of male-sterile cytoplasms in maize (Zea mays L.).
Crop Sci, 1971,11(5):724-727.
URL [本文引用: 1]
Thirty lines with male-sterile cytoplasm were backcrossed to a series of maize (Zea mays L.) inbreds. Each cytoplasm gave male-sterile plants in certain inbred backgrounds and partially or completely male-fertile plants in other backgrounds. The cytoplasms were divided into groups according to the pattern of fertility restoration observed. Each group appears to be composed of similar if not identical cytoplasms. The major groups are designated the Texas Group, the S Group, and the C Group. A fourth group (P) probably belongs to the Texas Group but may be distinct. A few sources of cytoplasm require further testing before they can be assigned to a group. C cytoplasm is assigned the permanent genetic symbol cms3.

[3] Zheng YL . Study on the mechanism of the fertility about several types of cytoplasmic male-sterility in maize
J Huazhong Agric College, 1982, ( 1):44-68.
[本文引用: 1]

郑用琏 . 若干玉米细胞质雄性不育类型(CMS)育性机理的研究
华中农学院学报, 1982, ( 1):44-68.
[本文引用: 1]

[4] Scheifele GL, Nelson RR, Koons C . Male sterility cytoplasm conditioning susceptibility of resistant inbred lines of maize to yellow leaf blight caused by Phyllosticta zeae
Plant Dis Rep, 1969,53:656-659.
URL [本文引用: 1]
Cytoplasmic control of susceptibility in maize to yellow leaf blight caused by Phyllostica zeae has been hypothesized in light of: (i) increased susceptibility to the disease of all inbreds with cytoplasm of the Texas male-sterile type, in contrast to the resistance of their normal counterparts; and (ii) uniformity in susceptibility of plants within the population of any inbred line with the Texas male-sterility type cytoplasm. Susceptibility largely governed by a cytoplasmic factor(s) conferring malesterility would insure uniform susceptibility to yellow leaf blight of maize hybrids, even though many inbred lines with a high degree of resistance are available.

[5] Feng Y, Zheng Q, Song H, Wang Y, Wang H, Jiang LJ, Yan JB, Zheng YL, Yue B . Multiple loci not only Rf3 involved in the restoration ability of pollen fertility, anther exsertion and pollen shedding to S type cytoplasmic male sterile in maize.
Theor Appl Genet, 2015,128(11):2341-2350.
URLPMID:26220224 [本文引用: 2]
Key message Thirty loci for fertility restoration of pollen fertility, anther exsertion and pollen shedding to maize CMS-S were identified by GWAS.

[6] 侯玮, 陈举林, 王国胜, 闫保罗, 李平海 . 细胞质雄性不育在玉米育种及生产中的应用概述
安徽农学通报, 2011,17(1):64-66.
URL [本文引用: 1]
玉米雄性不育系利用不仅是玉米育种技术的一项重大变革,也是种子 产业发展的必然趋势.通过分析细胞质雄性不育在玉米育种及生产中的应用现状,得出采用雄性不育技术,创造出具有国际竞争力的玉米新种质和新品种,提高国内 玉米种子生产质量,增强我国种子产业在国际市场上的竞争力,对于我国的种子产业的发展将会产生深远的影响.

[7] Sun QQ, Rong TZ . Study and use of cytoplasmic male sterility in maize
J Sichuan Agric Univ, 2003,21(1):49-53.
[本文引用: 1]

孙庆泉, 荣廷昭 . 玉米胞质雄性不育材料的研究和利用
四川农业大学学报, 2003,21(1):49-53.
[本文引用: 1]

[8] 李继耕 . 细胞质雄性不育性的分子机理
遗传, 1992,14(2):37-40.
URLMagsci [本文引用: 1]

[9] Li DD, Wang B . Rice atpA gene cloning and relationship with cutoplasimic male sterility.
Hereditas (Beijing), 1990,12(4):1-4.
URLMagsci [本文引用: 2]
本研究以水稻BT型细胞质雄性不育系秋光和相应的保持系秋光为材料，提取线粒体DNA，用限制性内切酶完全酶解，以玉米线粒体atpA基因和波菜叶绿体atpA基因作为探针，进行分子杂交，将保持系线粒体atpA基因定位在3.5kb的Bam HI酶切片段上，并且以pBR322为载体，克隆了这一片段，另外，在Bam HI完全酶解普带的杂交结果中，不育系线粒体基因组中有两条阳性杂交带，分别是3.5kb和2.9kb，而保持系线粒体基因组中只有3.5kb一条阳性杂交带，因而认为水稻不育系线粒体基因组中可能有两个atpA基因拷贝，而相应的保持系线粒体基因组中只有一个atpA基因拷贝。
李大东, 王斌 . 水稻线粒体atpA基因的克隆及其与细胞质雄性不育的关系
遗传, 1990,12(4):1-4.
URLMagsci [本文引用: 2]
本研究以水稻BT型细胞质雄性不育系秋光和相应的保持系秋光为材料，提取线粒体DNA，用限制性内切酶完全酶解，以玉米线粒体atpA基因和波菜叶绿体atpA基因作为探针，进行分子杂交，将保持系线粒体atpA基因定位在3.5kb的Bam HI酶切片段上，并且以pBR322为载体，克隆了这一片段，另外，在Bam HI完全酶解普带的杂交结果中，不育系线粒体基因组中有两条阳性杂交带，分别是3.5kb和2.9kb，而保持系线粒体基因组中只有3.5kb一条阳性杂交带，因而认为水稻不育系线粒体基因组中可能有两个atpA基因拷贝，而相应的保持系线粒体基因组中只有一个atpA基因拷贝。

[10] Li XM, Zheng YL, Zhang FD, Zhu YG . RFLP analysis for mitochondria genome of CMS rice Honglian type
Hereditas (Beijing), 2000,22(4):201-204.
URLMagsci [本文引用: 1]
应用ＲＦＬＰ技术，研究了红莲型不育系、保持系、杂种一代及野败型、马协型不育系的线粒体基因。结果表明，红莲型育系与保持系线粒体基因组之间在多外基因区域存在明显差异，为红莲型细胞质雄性不育分子机理研究提供了线索；红莲型不育系与野败型不育系的线粒体基因组之间存在显著差异，马协型不育系与野败型不育系的线粒体基因组之间存在一定差异，在分子水平揭示了不育胞质的多样性。
李小明, 郑用琏, 张方东, 朱英国 . 红莲型细胞质雄性不育水稻线粒体DNA的RFLP分析
遗传, 2000,22(4):201-204.
URLMagsci [本文引用: 1]
应用ＲＦＬＰ技术，研究了红莲型不育系、保持系、杂种一代及野败型、马协型不育系的线粒体基因。结果表明，红莲型育系与保持系线粒体基因组之间在多外基因区域存在明显差异，为红莲型细胞质雄性不育分子机理研究提供了线索；红莲型不育系与野败型不育系的线粒体基因组之间存在显著差异，马协型不育系与野败型不育系的线粒体基因组之间存在一定差异，在分子水平揭示了不育胞质的多样性。

[11] Dewey RE, Timothy DH, Levings III CS . Chimeric mitochondrial genes expressed in the C male-sterile cytoplasm of maize
Curr Genet, 1991,20(6):475-482.
URLPMID:1664299 [本文引用: 1]
Aberrant recombinations involving the mitochondrial atp9, atp6 and coxII genes have created unique chimeric sequences in the C male0sterile cytoplasm ( cms-C ) of maize. An apparent consequence of the rearrangements is the interchanging of transcriptional and/or translational regulatory signals for these genes, and alterations in the reading frames encoding the atp6 and coxII genes in the C cytoplasm. Particularly unusual is the organization of the atp6 gene in cms-C mitochondria, designated atp6-C . The atp6-C sequence is a triple gene fusion product comprised of DNAs derived from atp9, atp6 and an open reading frame of unknown origin. Although there is no direct evidence indicating that these chimeric genes are responsible for the cytoplasmic male sterility (cms) trait, their novel arrangements and the strong correlation between these genes and the C type of male sterility suggest such a role.

[12] Luo DP, Xu H, Liu ZL, Guo JX, Li HY, Chen LT, Fang C, Zhang QY, Bai M, Yao N, Wu H, Wu H, Ji CH, Zheng HQ, Chen YL, Ye S, Li XY, Zhao XC, Li RQ, Liu YG . A detrimental mitochondrial-nuclear interaction causes cytoplasmic male sterility in rice
Nat Genet, 2013,45(5):573-577.
URLPMID:23502780 [本文引用: 1]
Plant cytoplasmic male sterility (CMS) results from incompatibilities between the organellar and nuclear genomes and prevents self pollination, enabling hybrid crop breeding to increase yields. The Wild Abortive CMS (CMS-WA) has been exploited in the majority of 'three-line' hybrid rice production since the 1970s, but the molecular basis of this trait remains unknown. Here we report that a new mitochondrial gene, WA352, which originated recently in wild rice, confers CMS-WA because the protein it encodes interacts with the nuclear-encoded mitochondrial protein COX11. In CMS-WA lines, WA352 accumulates preferentially in the anther tapetum, thereby inhibiting COX11 function in peroxide metabolism and triggering premature tapetal programmed cell death and consequent pollen abortion. WA352-induced sterility can be suppressed by two restorer-of-fertility (Rf) genes, suggesting the existence of different mechanisms to counteract deleterious cytoplasmic factors. Thus, CMS-related cytoplasmic-nuclear incompatibility is driven by a detrimental interaction between a newly evolved mitochondrial gene and a conserved, essential nuclear gene.

[13] Kazama T, Toriyama K . A pentatricopeptide repeat-containing gene that promotes the processing of aberrant atp6 RNA of cytoplasmic male-sterile rice
FEBS Lett, 2003,544(1-3):99-102.
URLPMID:12782297 [本文引用: 1]
Abstract A fertility restorer gene (Rf-1) of [ms-bo] cytoplasmic male sterility (CMS) in rice has been reported to be responsible for the processing of RNA of aberrant atp6 of mitochondria. We have carried out map-based cloning of the Rf-1 gene and found that a 4.7-kb genomic fragment of a restorer line promoted the processing of aberrant atp6 RNA when introduced into a CMS line. The genomic fragment contained a single open reading frame encoding 18 repeats of the 35 amino acid pentatricopeptide repeat (PPR) motif. The cloned PPR gene is a possible candidate of Rf-1. A non-restoring genotype was identified to have deletions within the coding region.

[14] Krishnasamy S, Makaroff CA . Organ-specific reduction in the abundance of a mitochondrial protein accompanies fertility restoration in cytoplasmic male-sterile radish
Plant Mol Biol, 1994,26(3):935-946.
URLPMID:8000006
Abstract The mitochondrial DNA of plants containing the male sterility-causing Ogura cytoplasm of radish contain a novel gene, orf138, that is transcribed as part of a bicistronic mRNA. Genetic studies have previously linked male sterility with the orf138 locus. To determine if orf138 is expressed at the protein level, and investigate the effect of fertility restoration on ORF138 levels, we have raised antibodies to an ORF138-glutathione S-transferase fusion protein. Anti-ORF138 antibodies detect a 20 kDa protein that is associated with the mitochondrial membrane of sterile Ogura radish plants. Nuclear restoration is accompanied by a dramatic reduction in the amount of this protein in mitochondria of flowers and leaves, but not roots of fertile Ogura radish plants. The presence or absence of fertility restoration genes has no detectable effect on the size, abundance, or RNA editing patterns of orf138 transcripts. These results support genetic studies that have implicated orf138 in Ogura cytoplasmic male sterility and suggest that the restorer genes may be affecting either the translation or stability of ORF138.

[15] Liu Z, Dong FM, Wang X, Wang T, Su R, Hong DF, Yang GS . A pentatricopeptide repeat protein restores nap cytoplasmic male sterility in Brassica napus.
J Exp Bot, 2017,68(15):4115-4123.
URLPMID:28922764
react-text: 98 to idenify major QTls for important yield trais to construct genetic maps for these QTLs /react-text react-text: 99 /react-text

[16] Wei WL, Wang HZ, Liu GH . Advanced biological studies on the cytoplasmic male sterility and its fertility Restoration in plants
Hereditas (Beijing), 2005,27(4):651-658.
URLMagsci [本文引用: 1]
植物细胞质雄性不育性与育性恢复的分子机理一直是分子生物学的研究热点。文章综述了近十年来的主要研究进展。包括：1、线粒体不育相关区域的确定及其特点；2、不育相关区域的表达谱；3、产生细胞质雄性不育的可能机理；4、恢复基因的可能调控方式；5、恢复基因的遗传、定位及其特征等。拟兰芥、水稻等模式植物线粒体基因组测序工作已经完成，其有关生物学信息及后续研究将极大地推动植物细胞质雄性不育研究取得更快进展。
危文亮, 王汉中, 刘贵华 . 植物细胞质雄性不育性与育性恢复的分子生物学研究进展
遗传, 2005,27(4):651-658.
URLMagsci [本文引用: 1]
植物细胞质雄性不育性与育性恢复的分子机理一直是分子生物学的研究热点。文章综述了近十年来的主要研究进展。包括：1、线粒体不育相关区域的确定及其特点；2、不育相关区域的表达谱；3、产生细胞质雄性不育的可能机理；4、恢复基因的可能调控方式；5、恢复基因的遗传、定位及其特征等。拟兰芥、水稻等模式植物线粒体基因组测序工作已经完成，其有关生物学信息及后续研究将极大地推动植物细胞质雄性不育研究取得更快进展。

[17] Janska H, Sarria R, Woloszynska M, Arrieta-Montiel M, Mackenzie SA . Stoichiometric shifts in the common bean mitochondrial genome leading to male sterility and spontaneous reversion to fertility
Plant Cell, 1998,10(7):1163-1180.
URL [本文引用: 1]

[18] Mackenzie SA, Chase CD . Fertility restoration is associated with loss of a portion of the mitochondrial genome in cytoplasmic male-sterile common bean
Plant Cell, 1990,2(9):905-912.
URLPMID:12354971 [本文引用: 1]
Abstract Restoration of pollen fertility to cytoplasmic male-sterile common bean by nuclear gene Fr is accompanied by mitochondrial (mt) DNA rearrangements within restored plants. These rearrangements are also observed upon spontaneous cytoplasmic reversion to fertility. An mtDNA fragment of at least 25 kilobases was lost from the genome upon restoration or reversion. This fragment contained DNA segments that were not repeated elsewhere in the genome and, therefore, were not detected within the genome upon fertility restoration. This result suggested that the particular mtDNA configuration absent from restored plants could not be maintained by a constant process of recombination but rather by autonomous replication. No evidence of excision of this region from the mt genome, in the form of a junction fragment associating flanking DNA regions, was detected in fertile restored plants. DNA gel blot hybridization of this mtDNA region, compared with hybridization to related regions of the mitochondrial genome that shared sequence homology, indicated that the mtDNA region associated with sterility was present in lower copy number. These observations, as well as the occurrence of similar or identical rearrangements upon spontaneous cytoplasmic reversion, indicate that the restoration of pollen fertility may be accompanied by loss of an independently replicating subgenomic DNA molecule from the mitochondrial genome.

[19] He S, Lyznik A, Mackenzie S . Pollen fertility restoration by nuclear gene Fr in Cms bean: nuclear-directed alteration of a mitochondrial population.
Genetics, 1995,139(2):955-962.
URLPMID:7713444 [本文引用: 1]
Abstract Two nuclear genes, Fr and Fr2, have been identified that restore pollen fertility to cytoplasmic male sterile (CMS) common bean (Phaseolus vulgaris L.) by apparently distinct mechanisms. Whereas Fr2 appears to suppress the expression of a male sterility associated mitochondrial sequence (designated pvs), Fr restores pollen fertility by causing the elimination of this unusual mitochondrial DNA-segment. To further investigate the mechanism of Fr action, Fr and Fr2 were cointroduced into the nucleus of a bean line containing the sterility inducing cytoplasm. When the effect of pvs was suppressed by Fr2, the presence of Fr no longer directed the elimination of the mitochondrial pvs sequence. This result suggests that the Fr function is dependent on proper expression of the pvs sequence. To evaluate the temporal and spatial patterns of Fr action, we undertook a polymerase chain reaction-based approach to trace the fate of the pvs sequence in different tissues of F2 and F3 fertile-restored plants derived from a genetic cross between a cytoplasmic male sterile line of common bean, CMS-Sprite (frfr), and fertility restorer line R351 (FrFr). We demonstrate that the Fr-directed disappearance of pvs sequence occurs during flower development. Elimination of the pvs sequence from developing megaspores results in permanent fertility restoration in the following generations. Genetic analysis demonstrated that permanent fertility restoration, that is, the complete elimination of pvs from reproductive tissues requires two doses of the Fr allele or the absence of fr in F2 individuals.(ABSTRACT TRUNCATED AT 250 WORDS)

[20] Chen WC, Luo FH, Ji LY . Some genetic aspects of the c-type cytoplasmic male-sterility in maize and its use in breeding
Acta Agron Sin, 1979,5(4):21-28.
[本文引用: 2]

陈伟程, 罗福和, 季良越 . 玉米C型胞质雄花不育的遗传及其在生产上的应用
作物学报, 1979,5(4):21-28.
[本文引用: 2]

[21] Chen WC, Duan SF . The genetics of fertility C-cytoplasmic male sterility in corn
Acta Agric Univ Henanensis, 1986,20(2):125-140.
[本文引用: 1]

陈伟程, 段韶芬 . 玉米C型胞质雄性不育恢复性遗传研究
河南农业大学学报, 1986,20(2):125-140.
[本文引用: 1]

[22] Kheyr-Pour A, Gracen VE, Everett HL . Genetics of fertility restoration in the C-Group of cytoplasmic male sterility in maize
Genetics, 1981,98(2):379-388.
URLPMID:17249089 [本文引用: 1]
The genetics of fertility restoration of cms-C group cytoplasm of maize was studied using crosses involving stable maintainer lines and lines that restored full pollen fertility. Pollen fertility in the sources of cms-C sterile cytoplasms studied was restored by a single dominant restorer (Rf4) gene. The fertility restoration was sporophytic. Allelism tests among five restorer lines showed that they all apparently carried the same alleles (Rf4 Rf4). Similar tests also demonstrated that seven nonrestoring maintainer lines had apparently the same genotype (rf4 rf4), although a partial "late break" of fertility was observed at low levels in some maintainer crosses. Comparative studies among different cms-C sources (C, Bb, ES, PR and RB) indicated that similar inheritance of fertility restoration was involved. The data indicated that a single, dominant Rf gene is involved in the restoration of several C-group cytoplasms, at least in the lines studied here. This is the first single-gene, sporophytic restorer system described in maize to date.

[23] Sisco PH . Duplications complicate genetic mapping of Rf4, a restorer gene for cms-C cytoplasmic male sterility in corn.
Crop Sci, 1991,31(5):1263-1266.
URL [本文引用: 1]
Previous studies of the genetics of fertility restoration for C-type cytoplasmic male sterility (cms-C) in corn (Zea mays L.) have produced conflicting results. The objective of this study was to localize the restorer gene(s) for cms-C using restriction fragment length polymorphism (RFLP) analysis. Rf4, a single dominant restorer gene for cms-C found in inbred A619, was localized to Chromosome 8, almost-equal-to 2 centimorgans from an RFLP marker locus, NPI114A. The presence in corn of extensive nucleotide duplication complicated the analysis. Because the region of Chromosome 8 near Rf4 is duplicated on Chromosome 3, an additional restorer gene for cms-C may eventually be found on Chromosome 3.

[24] Hu YM, Tang JH, Yang H, Xie HL, Lu XM, Niu J H, Chen WC . Identification and mapping of Rf-I an inhibitor of the Rf5 restorer gene for Cms-C in maize( Zea mays L.).
Theor Appl Genet, 2006,113(2):357-360.
URLPMID:16791701 [本文引用: 2]
The restoration of the C-type cytoplasmic male sterility (Cms) has been a common agriculture practice in the production of hybrid seed for many years. In this study, a series of crosses between select sterile and restorer lines, as well as a backcross population of (Cms-C7702×026233)02×026233, were used to investigate the restoration of C-type Cms. Our results demonstrated that there was an inhibitor of the Rf5 restorer gene. This inhibitor gene, Rf-I , maps to chromosome 7 and is tightly linked with SSR markers, umc2326 and umc2327, at a genetic distance 4.7 and 3.402cM, respectively. After analyzing our data combined with previous studies, we propose that the restoration of C-type Cms has two dominant genes, Rf4 and Rf5 . Rf4 has the ability to restore all genotypes of Cms-C lines; however, there exists an inhibitor for the other restorer gene, Rf5 ; thus, it can restore only those genotypes of Cms-C lines lacking the Rf-I inhibitor.

[25] Liu ZH, Chen WC . Research on stability of male fertile restoration of corn hybrids with different sterile cytoplasms-II restorability relation to environments
Acta Agricul Univ Henanensis, 1992,26(1):1-8.
[本文引用: 1]

刘宗华, 陈伟程 . 玉米不育胞质杂交种雄花育性恢复稳定性的研究—Ⅱ恢复性与环境的关系
河南农业大学学报, 1992,26(1):1-8.
[本文引用: 1]

[26] Weider C, Stamp P, Christov N, Hüsken A, Foueillassar X, Camp K, Munsch M . Stability of cytoplasmic male sterility in maize under different environmental conditions
Crop Sci, 2009,49(1):77-84.
URL [本文引用: 1]
Cytoplasmic male sterility (CMS) is a maternally transmitted trait, whereby a plant is unable to produce viable pollen. Studies have revealed that this trait is a tool for enabling efficient and reliable coexistence between genetically modified (GM) and non-GM cultivation by biocontainment of GM maize (Zea mays L.) pollen. Maize has three types of male-sterile cytoplasm (T, S, and C), the fertility of which can be restored by nuclear rf genes or by interactions with the environment. Twenty-two CMS versions of modern European maize hybrids were evaluated in 17 environments in Switzerland, France, and Bulgaria, with two or three sowing dates, in 2005 and 2006. Stable and unstable male sterility occurred in all three CMS types. T-cytoplasm hybrids were the most stable, while S-cytoplasm hybrids often showed partial restoration of fertility. C-cytoplasm was similar to T-cytoplasm with regard to maintaining male sterility. Climatic factors, especially air temperature, evapotranspiration, and water vapor, during the 10 d before anthesis as well as during anthesis, were correlated positively or negatively with the partial reversion to male fertility of CMS hybrids, indicating an interaction between genetic and climatic factors. This study illustrates that T- and C-cytoplasm in particular open up viable prospects for containing transgenic pollen, especially for Bt-maize.

[27] Bueckmann H, Thiele K, Schiemann J . Influence of soil moisture and air temperature on the stability of cytoplasmic male sterility (CMS) in maize (Zea mays L.).
Agric Sci, 2016,7(2):70-81.
URL [本文引用: 1]
Cytoplasmic male sterility (CMS) is a maternally inherited trait that suppresses the production of viable pollen. CMS is a useful biological tool for confinement strategies to facilitate coexistence of genetically modified (GM) and non-GM crops in case where it is required. The trait is reversible and can be restored to fertility in the presence of nuclear restorer genes (Rf genes) and by environmental impacts. The aim of this study was to investigate the influence of the level of irrigation on the stability of CMS maize hybrids under defined greenhouse conditions. Additionally the combination of irrigation and air temperature was studied. Three CMS maize hybrids were grown with different levels of irrigation and in different temperature regimes. Tassel characteristics, pollen production and fertility were assessed. The CMS stability was high in hot air temperatures and decreased in lower temperatures. The level of irrigation had no major effect on the level of sterility. The extent of these phenomena was depending on the genotype of CMS maize and should be known before using CMS for coexistence purposes.

[28] Tracy WF, Everett HL, Gracen VE . Inheritance, environmental effects, and partial male fertility in C-type CMS in a maize inbred
J Hered, 1991,82(4):343-346.
URL [本文引用: 1]
ABSTRACT While late-breaking partial fertility restoration of cytoplasmic male sterility is important both economically and theoretically, its inheritance is not well understood. Sublines of the maize inbred A632 differing in the level of restoration of C-cytoplasmic male sterility were identified. One line was nearly completely male sterile, whereas the others had varying levels of partial restoration; full fertility restoration was not observed. We crossed the fertile selections to the male sterile one and evaluated the progeny of the crosses to demonstrate the genetic nature of these differences. In two different experiments the percentage of plants that exhibited partial fertility was significantly different ( P =.01). In experiment I, the progeny of the cross A632-Pr-LB-1 A632-Pr-MS were 55% partially fertile while the progeny of A632-Pr-LB-2 A632-Pr-MS were 75% partially fertile. In experiment II, 3% of the plants exhibited partial fertility in one cross while 9% did so in the other cross. All differences were significant. Progeny tests were performed over years, and the effect of years on fertility restoration was highly significant. The number of fertile individuals was strongly affected by the environment with only 1.4% of the plants shedding pollen in 1984 and 13.0% shedding pollen in 1985. However, there was no environment by inbred interaction for fertility restoration. Due to low penetrance and strong environmental effects it was not possible to estimate the number of genes involved. However, it appears that this trait is under polygenic control.

[29] Chen W, Liu ZX , E LZ, Yang H, Dai JR. Classification of male sterile cytoplasm of CMS-P in maize (
Zea mays everta). Acta Agron Sin, 2007,33(2):196-200.
Magsci [本文引用: 1]
<P>采用育性恢复专效性测定、PCR分类鉴定、类质粒鉴定3种玉米不育细胞质分类方法，对本室从爆裂玉米（<EM>Zea mays everta</EM>）种质中发现的玉米细胞质雄性不育材料“CMS-P”进行了胞质分类研究，一致表明CMS-P属于S型不育胞质。这为CMS-P的有效利用提供了理论依据。卡方测验表明，S组田间育性表现恢复的各组合花粉可染率并不完全符合1∶1的分离比例，CMS-P向可育方向偏移的程度更大。花粉可染率在不同测验系、不同胞质、不同环境间都表现出较大差异，胞质与核微效基因间的互作可能是影响育性稳定性的主要因素。</P>
陈伟, 刘占先, 鄂立柱, 杨会, 戴景瑞 . 玉米细胞质雄性不育材料CMS-P的胞质分类研究
作物学报, 2007,33(2):196-200.
Magsci [本文引用: 1]
<P>采用育性恢复专效性测定、PCR分类鉴定、类质粒鉴定3种玉米不育细胞质分类方法，对本室从爆裂玉米（<EM>Zea mays everta</EM>）种质中发现的玉米细胞质雄性不育材料“CMS-P”进行了胞质分类研究，一致表明CMS-P属于S型不育胞质。这为CMS-P的有效利用提供了理论依据。卡方测验表明，S组田间育性表现恢复的各组合花粉可染率并不完全符合1∶1的分离比例，CMS-P向可育方向偏移的程度更大。花粉可染率在不同测验系、不同胞质、不同环境间都表现出较大差异，胞质与核微效基因间的互作可能是影响育性稳定性的主要因素。</P>

[30] Yang GS, Fu TD . Environment effects on the cytoplasmic male-sterility of rapeseed (B. napus and B. campestris).
Chin J Oil Crop Sci, 1987, ( 3):15-19.
[本文引用: 1]

杨光圣, 傅廷栋 . 环境条件对油菜细胞质雄性不育的影响
中国油料作物学报, 1987, ( 3):15-19.
[本文引用: 1]

[31] Kohls S, Stamp P, Knaak C, Messmer R . QTL involved in the partial restoration of male fertility of C-type cytoplasmic male sterility in maize
Theor Appl Genet, 2011,123(2):327-338.
URLPMID:21479555 [本文引用: 2]
Partial restoration of male fertility limits the use of C-type cytoplasmic male sterility (C-CMS) for the production of hybrid seeds in maize. Nevertheless, the genetic basis of the trait is still unknown. Therefore, the aim to this study was to identify genomic regions that govern partial restoration by means of a QTL analysis carried out in an F 2 population ( n = 180). This population was derived from the Corn Belt inbred lines B37C and K55. F 2 BC 1 progenies were phenotyped at three locations in Switzerland. Male fertility was rated according to the quality and number of anthers as well as the anthesis-silking interval. A weak effect of environment on the expression of partial restoration was reflected by high heritabilities of all fertility-related traits. Partial restoration was inherited like an oligogenic trait. Three major QTL regions were found consistently across environments in the chromosomal bins 2.09, 3.06 and 7.03. Therefore, a marker-assisted counter-selection of partial restoration is promising. Minor QTL regions were found on chromosomes 3, 4, 5, 6 and 8. A combination of partial restorer alleles at different QTL can lead to full restoration of fertility. The maternal parent was clearly involved in the partial restoration, because the restorer alleles at QTL in bins 2.09, 6.04 and 7.03 originated from B37. The three major QTL regions collocated with other restorer genes of maize, a phenomenon, which seems to be typical for restorer genes. Therefore, a study of the clusters of restorer genes in maize could lead to a better understanding of their evolution and function. In this respect, the long arm of chromosome 2 is particularly interesting, because it harbors restorer genes for the three major CMS systems (C, T and S) of maize.

[32] Duvick DN . Cytoplasmic pollen sterility in corn
Adv Genet, 1965,13:1-56.
URL [本文引用: 2]
This chapter discusses the general field of cytoplasmic inheritance of pollen sterility. In five discoveries, the expression of the cytoplasmically induced sterility was influenced by nuclear genes. Investigation of the genetics of fertility restoration of both the United States Department of Agriculture (USDA) and the Texas types of cytoplasmic sterility has shown that in either case a single dominant gene is sufficient to restore pollen fertility to most cytoplasmic male-sterile inbreds although there are complications with certain genetic backgrounds and in certain environments. Specific tester genotypes enable one to differentiate USDA from Texas cytoplasms because some genotypes are sterile in one cytoplasm but not the other, some are sterile in both, and some are pollen fertile in both. This type of test can be applied to all discoveries of cytoplasmic male sterility to see if they resemble either the USDA or the Texas source or if they have a different spectrum of interactions with genotype. Cytoplasmic male sterility in corn is characterized by having two distinct forms, the USDA and the Texas. Both types interact with the nucleus to the extent that their sterility-inducing potential is revealed only by certain genes, and the number of the genes is relatively few for each cytoplasm. The Texas type has been shown to affect, to a small degree, the rate of growth and final size of the corn plant, independently of its effect on pollen sterility.

[33] Zheng YL, Liu JL . Relativity between mitochondria DNA and fertility instability of CMS-S in maize
J Huazhong Agric Univ, 1992,11(2):120-126.
[本文引用: 1]

郑用琏, 刘纪麟 . 玉米CMS-C材料雄花育性不稳定性与DNA的关系
华中农业大学学报, 1992,11(2):120-126.
[本文引用: 1]

[34] Chen WC, Duan SF . The cytological observation of anther development in C-cytoplasmic male-sterile corn
Acta Agron Sin, 1988,14(3):177-181.
Magsci [本文引用: 1]
对3个 C 型不育系及其同型保持系和4个恢复系与这3个不育系组配的12个 F_1的花药发育过程进行了观察,以了解 C 型不育系雄性败育的细胞学机理。观察结果表明,不同核背景的不育系,其雄性败育的细胞学特点各异。Cms-C Mo17的花药绒毡层细胞为双核型。该层细胞在减数分裂的四分体时期开始液泡化,并呈辐射状膨大,至小孢子的单核前
陈伟程, 段韶芬 . 玉米C型胞质雄性不育系花药发育的细胞学观察
作物学报, 1988,14(3):177-181.
Magsci [本文引用: 1]
对3个 C 型不育系及其同型保持系和4个恢复系与这3个不育系组配的12个 F_1的花药发育过程进行了观察,以了解 C 型不育系雄性败育的细胞学机理。观察结果表明,不同核背景的不育系,其雄性败育的细胞学特点各异。Cms-C Mo17的花药绒毡层细胞为双核型。该层细胞在减数分裂的四分体时期开始液泡化,并呈辐射状膨大,至小孢子的单核前

[35] Zhai HQ . A hypothesis of threshold character and dosage effector the hereditary manners of male sterile and fertility restoring genes in rice
Hereditas (Beijing), 1990,12(5):47-48.
URLMagsci [本文引用: 2]
杂交水稻不育一恢复基因的遗传行为比较复杂，现有的关于该方面研究的报道不足以解释杂交水稻不育一恢复基因的遗传机理。本文提出杂交水稻不育一恢复基因的阑性状和剂量效应假说，其主要论点有二：(1)恢复基因只有剂量效应，没有显隐性之分，一个品种恢复力的强弱依赖于该品种携带恢复基因的剂量；(2）不育一恢复这一性状属于闻性状，是核质互作的结果，当合子中恢复基因的剂量达到某rA值时，激活了不育系细胞质的育性，因此对于某一不育系来说，其相应的恢保关系可以分为三大类：没有恢复能力的L型品种，可作为保持系；具有完全恢复能力的H型品种，可以作为恢复系；具有部分恢复能力的M型品种。根据此假说得到的推论能够解释杂交水稻不育一恢复基因的遗传行为。
翟虎渠 . 水稻雄性不育—恢复基因的阈性状和剂量效应假说
遗传, 1990,12(5):47-48.
URLMagsci [本文引用: 2]
杂交水稻不育一恢复基因的遗传行为比较复杂，现有的关于该方面研究的报道不足以解释杂交水稻不育一恢复基因的遗传机理。本文提出杂交水稻不育一恢复基因的阑性状和剂量效应假说，其主要论点有二：(1)恢复基因只有剂量效应，没有显隐性之分，一个品种恢复力的强弱依赖于该品种携带恢复基因的剂量；(2）不育一恢复这一性状属于闻性状，是核质互作的结果，当合子中恢复基因的剂量达到某rA值时，激活了不育系细胞质的育性，因此对于某一不育系来说，其相应的恢保关系可以分为三大类：没有恢复能力的L型品种，可作为保持系；具有完全恢复能力的H型品种，可以作为恢复系；具有部分恢复能力的M型品种。根据此假说得到的推论能够解释杂交水稻不育一恢复基因的遗传行为。

[36] Geyer M, Albrecht T, Hartl L, Mohler V . Exploring the genetics of fertility restoration controlled by Rf1 in common wheat( Triticum aestivum L.) using high-density linkage maps.
Mol Genet Genomics, 2018,293(2):451-462.
URLPMID:29177892 [本文引用: 1]
Abstract Hybrid wheat breeding has the potential to significantly increase wheat productivity compared to line breeding. The induction of male sterility by the cytoplasm of Triticum timopheevii Zhuk. is a widely discussed approach to ensure cross-pollination between parental inbred lines in hybrid wheat seed production. As fertility restoration in hybrids with this cytoplasm is often incomplete, understanding the underlying genetics is a prerequisite to apply this technology. A promising component for fertility restoration is the restorer locus Rf1, which was first detected on chromosome 1A of the restorer accession R3. In the present study, we performed quantitative trait locus (QTL) analyses to locate Rf1 and estimate its effect in populations involving the restorer lines R3, R113 and L19. Molecular markers linked to Rf1 in these populations were used to analyse the genomic target region in T. timopheevii accessions and common wheat breeding lines. The QTL analyses revealed that Rf1 interacted with a modifier locus on chromosome 1BS and the restorer locus Rf4 on chromosome 6B. The modifier locus significantly influenced both the penetrance and expressivity of Rf1. Whereas Rf1 exhibited expressivity higher than that of Rf4, the effects of these loci were not additive. Evaluating the marker haplotype for the Rf1 region, we propose that the restoring Rf1 allele may be derived exclusively from T. timopheevii. The present study demonstrates that interactions between restorer and modifier loci play a critical role in fertility restoration of common wheat with the cytoplasm of T. timopheevii.

[37] Yu SW, Xie JK, Wan Y, Hu BL . Genetic study and environmental effects on fertility restoration of cytoplasmic male sterility of rice
Acta Agric Jiangxi, 2004,16(4):5-9.
[本文引用: 1]

余守武, 谢建坤, 万勇, 胡标林 . 水稻细胞质雄性不育育性恢复遗传及其环境效应研究
江西农业学报, 2004,16(4):5-9.
[本文引用: 1]

[38] Nico DS, Danny G . The impact of environmental stress on male reproductive development in plants: biological processes and molecular mechanisms
Plant Cell Environ, 2014,37(1):1-18.
URLPMID:4280902 [本文引用: 1]
Abstract In plants, male reproductive development is extremely sensitive to adverse climatic environments and (a)biotic stress. Upon exposure to stress, male gametophytic organs often show morphological, structural and metabolic alterations that typically lead to meiotic defects or premature spore abortion and male reproductive sterility. Depending on the type of stress involved (e.g. heat, cold, drought) and the duration of stress exposure, the underlying cellular defect is highly variable and either involves cytoskeletal alterations, tapetal irregularities, altered sugar utilization, aberrations in auxin metabolism, accumulation of reactive oxygen species (ROS; oxidative stress) or the ectopic induction of programmed cell death (PCD). In this review, we present the critically stress-sensitive stages of male sporogenesis (meiosis) and male gametogenesis (microspore development), and discuss the corresponding biological processes involved and the resulting alterations in male reproduction. In addition, this review also provides insights into the molecular and/or hormonal regulation of the environmental stress sensitivity of male reproduction and outlines putative interaction(s) between the different processes involved. 2013 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.