‘新高’梨花粉败育的细胞学和生理特性研究
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摘要
梨(Pyrus)属于典型的自交不亲和性植物，生产中需要合理配置授粉树，才能达到预期的产量和品质。在栽培的品种中，一些雄性败育的品种，如‘新高’、‘爱宕’和‘黄金梨’等，增加了授粉树选配的问题和生产管理上的困难。为克服梨雄性不育给生产上带来的负面影响和利用雄性不育资源在解决梨花粉直感效应及改善品质方面的潜在价值，本试验以雄性不育‘新高’梨和雄性可育‘丰水’梨为材料，开展梨雄性不育机理的研究。为此，本试验在对‘新高’梨雄性不育性进行鉴定的基础上，研究其花粉败育的时期和特征；并以此为依据，从细胞和亚细胞水平以及荧光物质的分布特性等方面研究了‘新高’花粉败育的细胞学机制；分析花蕾发育过程中矿质营养变化，对Ca~(2+)及其影响Ca~(2+)分布的酶进行了细胞化学定位，以探讨其分布的变化与花粉败育的关系；最后对花蕾中的激素及其影响IAA的酶和多胺进行分析，以从生理角度进一步揭示‘新高’花粉败育的机制。研究主要结果如下：
1．‘新高’花药皱缩，发白，虽可以正常开裂，但花粉很少。在少数花药的横切面上虽然可看到微量花粉，但花粉染色很浅，且不规则，没有生活力。‘新高’花粉母细胞能正常进行减数分裂产生四分体；在小孢子产生后，发育缓慢，小孢子壁薄，加厚不明显；随着单核小孢子的液泡化，小孢子粘连降解，或小孢子细胞壁破损，细胞质、细胞核降解消失，很少见到二核花粉。说明‘新高’属于小孢子退化类型的雄性不育品种，小孢子退化主要发生在单核小孢子中期到晚期。
2．在花粉母细胞减数分裂前，‘新高’花粉母细胞和花药壁的发育与‘丰水’没差异，药壁包含表皮、药室内壁、中层和绒毡层，花粉母细胞被胼胝质包围。‘新高’四分小孢子染色很浅，胼胝质解体后，‘新高’较‘丰水’小孢子浅染，外粉壁发育迟缓而显得较薄。单核中期大部分小孢子收缩变形，且大小不等，细胞质解体，大部分不见细胞核。随后，小孢子壁破损，内含物外流，小孢子逐渐解体成为空壳或仅留下小孢子壁降解的残余物。在游离小孢子形成时，‘新高’绒毡层出现了不均匀的染色，其它花药壁结构与‘丰水’差异不明显。
3．‘新高’在小孢子母细胞时期出现包裹小孢子母细胞的胼胝质厚薄不均匀，部分不完整，胞内线粒体等细胞器的结构模糊，内质网分布少等早期异常现象；在四分体小孢子内小囊泡分布少，线粒体等细胞器的膜呈透明状，之后小孢子液泡膜破裂，细胞质开始解体，线粒体、质体、内质网等细胞器相继解体或退化，核仁解体，最后出现了同心环状的膜状体，以及细胞质及细胞核降解的残骸。而‘丰水’小孢子母细胞和雄配子体的线粒体结构清晰，液泡膜完整，且含有丰富的内质网、核糖体和高尔基体等细胞器，此外在四分体小孢子膜内分散着大量的小囊泡，这些小泡可能与原外壁的形成有关。
4．‘新高’四分小孢子表现出局部形成原外壁，或质膜和原外壁分离等不良现象，同时在胞内有很少的小囊泡。在单核小孢子时期，外壁上不能沉积孢粉素或只有表面沉积孢粉素，形成了透明状的外壁或无基足层的外壁，即使有的可以形成完整的外壁，但较正常孢子的外壁薄。而‘丰水’在四分小孢子时期形成了良好的原外壁，在雄配子体发育过程中，其外壁迅速发育。在花粉成熟时，花粉壁由薄的内壁及由基足层、基柱、覆盖层共同组成的外壁构成。两个品种的小孢子母细胞或四分体时期的胼胝质均能正常形成，并能及时降解，没有差异；‘丰水’小孢子外壁的自发荧光物质随雄配子体的发育而逐渐增多，在单核小孢子后期，小孢子外壁上除萌发的孔沟外均匀地沉积大量的自发荧光物质。‘新高’在单核小孢子早期小孢子外壁的荧光物质与‘丰水’差异不大，之后从局部到整个小孢子表面呈透明状，随之变形和相互粘连，有的小孢子表面荧光物质呈网状分布等异常表现。
5．‘新高’和‘丰水’小孢子母细胞时期的绒毡层细胞都含有大量的线粒体、内质网等细胞器；随着减数分裂的进行，两个品种的绒毡层细胞质收缩，细胞间和内切相面形成了许多空腔，细胞内高尔基体和分泌小泡增多，内质网膨胀形成槽库结构；四分体时期，‘丰水’绒毡层内分泌小囊泡和槽库结构的内质网进一步增多，并向药室分泌大量的乌氏体，而‘新高’绒毡层细胞内的部分细胞器消失，内质网垛叠，线粒体等细胞器边缘呈透明状，并产生了大量的小液泡；在小孢子发育的过程中，‘新高’并没有像‘丰水’一样出现结构清晰的内质网、线粒体等细胞器，而且出现了一些形态各异、结构模糊的细胞器；在绒毡层细胞解体时，‘丰水’绒毡层细胞内主要为油体等结构，并呈极性分布。而‘新高’细胞内还含有槽库结构的内质网，并和其它结构一起呈无序分布。‘新高’和‘丰水’绒毡层中自发荧光物质都产生于小孢子母细胞时期，以后荧光增强。在四分体小孢子即将释放单核小孢子时，‘丰水’绒毡层细胞表面分布大量的自发荧光物质颗粒，而‘新高’表面的自发荧光物质颗粒较少。
6．‘新高’结果枝和结果母枝中Fe、Zn、Cu、Ca、Mg和K含量较‘丰水’低，尤其是结果枝中ca和zn的含量差异更大，只有Mn的差异不显著。从四分体时期开始到单核小孢子后期，‘新高’和‘丰水’梨在花蕾中各矿质营养含量总体上呈逐渐下降的趋势，但‘新高’中Fe、zn、cu、Mg和Ca含量均低于‘丰水’，差异达到显著水平，Mn含量在单核花粉期后显著地高于‘丰水’，K的差异不明显。
7．运用焦锑酸钾沉淀法，对‘新高’和‘丰水’花药发育过程中Ca~(2+)进行了细胞化学定位研究。‘新高’和‘丰水’小孢子母细胞膜表面均有Ca~(2+)淀颗粒的分布。在四分体时期，‘丰水’小孢子表面积累了大量的Ca~(2+)沉淀颗粒，而在‘新高’中基本没有Ca~(2+)沉淀颗粒。随着胼胝质的解体，在‘丰水’小孢子的液泡、外壁以及细胞膜等部位均积累了Ca~(2+)沉淀颗粒，‘新高’小孢子中的Ca~(2+)沉淀颗粒则更多地分布在细胞质以及一些细胞器上。‘新高’小孢子中的Ca~(2+)常分布与其花粉败育有关。从小孢子母细胞开始，‘丰水’花药绒毡层的径切向面和内切向面均分布着大小不等的Ca~(2+)沉淀颗粒。随着小孢子的产生和发育，Ca~(2+)沉淀颗粒分布的量逐渐增多。在单核晚期，Ca~(2+)沉淀颗粒又减少。但‘新高’在四分体时期绒毡层中Ca~(2+)沉淀颗粒明显比‘丰水’要少，而单核小孢子时期Ca~(2+)分布较‘丰水’多。
8．应用铅沉淀法，分析了花药发育过程中Ca~(2+)-ATPase的分布。‘丰水’小孢子母细胞膜上基本上没有Ca~(2+)-ATPase的分布，只是在一些细胞器上有少量的分布；单核小孢子发育过程中，‘丰水’小孢子质膜及内壁上的Ca~(2+)-ATPaS逐渐增加，但在胞质内分布较少。从小孢子母细胞时期开始，在‘丰水’绒毡层细胞膜和液泡膜等细胞器以及乌氏体上分布有大量的Ca~(2+)-ATPase，随绒毡层的解体而逐渐减少。自四分小孢子开始，‘新高’和‘丰水’花药Ca~(2+)-ATPase就有差别：‘新高’四分小孢子细胞膜几乎没有Ca~(2+)-ATPase的分布，随小孢子的发育质膜上Ca~(2+)-ATPase虽逐渐增多，但比‘丰水’明显少；‘新高’绒毡层膜以及乌氏体有明显Ca~(2+)-ATPase分布，除小孢子母细胞时期外，与‘丰水’的差异不明显。
9．‘新高’梨花蕾中IAA含量从四分体后极显著地低于‘丰水’，在单核花粉早期和单核花粉晚期仅为‘丰水’的22．7％和27．0％，‘新高’梨IAA含量的亏缺可能是导致‘新高’败育的原因之一；‘新高’梨在花粉发育的前期IAA氧化酶活性显著地高于‘丰水’，而两个品种POD活性变化一致，并且活性的差异不显著，说明POD不是导致‘新高’IAA含量的亏缺的关键酶；‘新高’梨ABA含量从四分体以后却显著地高于‘丰水’。
10．‘新高’花蕾发育过程中游离态的Put和Spd变化幅度不大，‘丰水’则在单核花粉到二核花粉时期出现高峰，其含量是‘新高’的3-5倍；‘新高’中游离态的Spm也显著地低于‘丰水’。‘丰水’中高氯酸可溶共价结合态Put、Spd和Spm含量逐渐增加，‘新高’在花粉败育期或败育前期出现不同于‘丰水’的高峰，此种形态多胺含量的增加可能影响到游离态多胺含量的增加，而影响到花粉的发育。‘新高’和‘丰水’中高氯酸不可溶结合态多胺含量的变化基本一致。‘丰水’中不同种类的多胺均以游离态为主，‘新高’以高氯酸可溶结合态为主。
综上所述，本研究从细胞形态、亚细胞水平、细胞化学和生理生化水平上首次对‘新高’雄性不育机理进行了系统的研究。明确了‘新高’花粉败育的时期和主要细胞学机制，探明了‘新高’雄性不育在细胞化学和某些生理生化方面的败育机理。为从分子水平来研究‘新高’雄性不育的机理和人工调控育性的表达提供了理论依据。
Pear (Pyrus) belongs to gametophytic self-incompatibility, and allogamy among somevarieties also occur incompatibility. So, the suitable pollination trees and artificalsupplementary pollination are employed to meet expectant yield and quality. Some malesterile varieties of pear in production, for example 'Niitaka', 'Atagi', 'whangkeumbae', andso on, increase difficulties in choice of pollination tree and management. In order toconquer negative influence of male sterile cultivars, utilize potential cast of male sterileresource to solve pollen xenia effect and improve fruit quality, male sterile mechanism ofpear was investigated with male sterile 'Niitake' and male fertile 'Housui' (Pyrus pyrifolaiNakai). In the present study, transmission electron microscope, fluorescent and opticalmicroscope and some modem analytic equipment were employed, and the technology offluorescence labeling and cyto-chemistry was introduced. Based on identification of malesterility of 'Niitaka', pollen abortive stage and cytological characteristics, and distributionof fluorescent material were studied during pollen abortion. Moreover, Ca~(2+) andCa~(2+)-ATPase in anther were located to indentify the relationship between their distributionand pollen abortion. In addtion, hormones and polyamines in floral bud were analyzed tofurther study physiological mechanism of male sterility in pear. Main results of presentstudy are as follows:
1. Anther of 'Niitaka' appeared shrinkage and whitening, and could normally take placedehiscence, but had a small quantity of pollen. There was a lack of pollen on transversesection of anther, or a few pollens were found on a few sections, but these pollens appearedlight dyeing, abnormal shape and lower germination rate. The microspore mother cell couldnormally occur meiosis and produce tetrad; Development of microspore was slow, and wallof microspore was thin and slowly thicken after microsporogenesis; Microspore adhered toeach other, or wall, cytoplasm and nucleus of microspore was disintegrated with microspore vacuolization, and two-called pollen was not observed. It was suggested that 'Niitaka'belong to male sterile cultivar of microspore degenerated form, pollen abortion of 'Niitaka'appeared from middle unimicrospore stage to late unimicrospore stage.
2. Technology of paraffin section was used to observe anther development. There was notdifference of microspore mother and anther wall of 'Niitaka' and 'Housui' before meiosis,and anther wall were composed of epidermis, endothecium, middle layer and tapetum,microspore mother cell was surrounded by callose wall. Dyeing of microspore of 'Niitaka'was very light in tetrad, and was still light after callose wall disintegrated. Wall ofmicrospore slowly thickened and was very thin. Most microspores shnmk, deformed andappeared different size, cytoplasm was disintegrated, and nucleus disappeared. Microsporebecame hollow shell or debris with wall damaged and inclusion spilled. Tapetal cellappeared irregular dyeing at the stage of microspore just separated from tetrad, and othercomposition of anther wall disappeared difference between 'Niitaka' and 'Housui'.
3. Microsporocyte and male gametophyte of 'Housui' had perfect mitochondria,integrated vacuole membrane and rich endoplasmic reticulum, ribosome and Golgi bodyunder transmission electron microscope. There were abundant vesicles in microspore attetrad. Callose wall which surrounded microsporocyte of 'Niitake' appeared irregulardistribution, microsporocyte showed abnormal appearance, for example indiscerniblemitochondria structure, a small quantity of endoplasmic reticulum and so on; There were afew vesicle and some organdies of transparent membrane like mitochondria in microsporeof tetrad. Cytoplasm begun to disintegrate with vacuole membrane damaged, followingmitochondria, plastid and endoplasmic reticulum disintegrated, and nucleolus wasdegenerated. At last, there were some membrane bodies of concentric ring and debris ofcytoplasm and nucleus degenerated in microspore.
4. Microspore of 'Housui' could form perfect primexine at the tetrad stage, and couldrapid produce exine with male gametophyte development. Wall of mature pollen werecomposed of thin intine and exine including foot layer, baculum and tectum. Microspore of'Niitaka' appeared a portion of primexine or primexine separated from membrane at thetetrad stage, and there were a small quantity of vesicle. Microspore showed lacksporopollenin in exine or a little sporopollenin on the surface of exine, so exine oftransparency or lack foot layer was formed. Although some microspores could formintegrated exine, exine was thinner than that of normal microspore. Two cultivars disappeared difference in callose wall of formation and disintegration, which surroundedmicrosporocyte and microspore at tetrad stage. Auto-fluorescent material of microsporewall gradually increased with male gametophyte development in 'Housui', andauto-fluorescent material homogeneously distributed on wall of microspore except forcolporate at late microspore stage. There was not obvious difference in auto-fluorescentmaterial on wall of microspore between 'Housui' and 'Niitaka' at early microspore stage,microspore surface appeared transparency from portion to whole with auto-fluorescentmaterial decrease, and microspore deformed and adhered to each other, auto-fluorescentmaterial showed reticular distribution on the surface of some microspore which had beendisintegrated.
5. Tapetal cell of 'Niitaka' and 'Housui' had abundant mitochondria and endoplasmicreticulum at microsporocyte stage; cytoplasm of tapetal cell shrunk with rnicrosporocytemeiosis of 'Niitaka' and 'Housui', lots of cavity was formed in intercellular space and innersurface of tapetum with tapetal cell shrinkage. Golgi body and vesicle gradually increasedin tapetal cell, endoplasmic reticulum became dilated cistamae. The number of excretivevesicle and endoplasmic reticulum with cistarae further increased in tapetal cell of 'Housui',and tapetal cell excreted lots of ubish body for locule at tetrad stage. There was someabnormal appearance in tapetal cell of 'Niitaka' at tetrad stage, including part organellesdegenerated, stacked endoplasmic reticulum, mitochondria of transparent edge and a lot ofvacuoles. Tapetal cell of 'Niitaka' did not have perfect mitochondria and endoplasmicreticulum as tapetal cell of 'Housui' had, and had deformed and indiscernible organelleswith microspore vacuolization; lipid body appeared polar distribution in tapetal cell of'Housui' when tapetal cell disintegrated. Endoplasmic reticulum with cistarae and othermaterial appeared irregular distribution in tapetal cell of 'Niitaka'. Auto-fluorescentmaterial of tapetum appeared at microsporocyte, and fluorescent density strengthened withtapetal development in two cultivars. A mass of auto-fluorescent material which wasaccumulated on the surface of tapetal cell of 'Housui' was released and regularly distributedon the surface of microspore. But auto-fluorescent material of 'Niitaka' was less than thatof 'Housui', irregularly distributed on the surface of microspore.
6. The contents of iron (Fe), zinc (Zn), copper (Cu), calcium (Ca), magnesium (Mg) andpotassium (K) in basal bearing shoot and bearing shoot of 'Niitaka' were lower than thoseof 'Housui', especially the differences of the content of Ca and Zn in bearing shoot were even more between two cultivars, but the differences of the manganese (Mn) content wasnot significant. Mineral nutrition content in floral bud of 'Niitaka' and 'Housui' appeareddescendant trend from tetrad stage to late uninucleate microspore stage, and the contents ofFe、Zn、Cu、Mg and Ca of 'Niitaka' was significantly lower than those of'Housui', the Mncontent was significantly higher than those of 'Housui, the difference of K content was notsignificant.
7. Potassium antimontate was used to locate Ca~(2+) in 'Niitaka' and 'Housui' anther. Theamount and distribution of calcium precipitates changed during pollen development andwere different between 'Niitaka' and 'Housui'. The calcium precipitates accumulated on thesurface of microsporocyte of 'Niitaka' and 'Housui'. Lots of calcium precipitates appearedon the surface of microspore of 'Housui' at tetrad stage, but did not appeared on the surfaceof microspore of 'Niitaka'. The calcium precipitates gradually accumulated on thecytoplasmic membrane, vacuole membrane and exine of microspore of 'Housui' aftercallose wall disintegrated, but the most calcium precipitates were distributed in cytoplasmand some organelles of microspore of 'Niitaka'. The calcium precipitates appeared on thesurface of tapetal cell of 'Housui' at microsporocyte stage, and gradually increased withmicrosporogenesis and development, decreased at late uninucleate microspore stage. Thecalcium precipitates on the surface of tapetal cell of 'Niitaka' were less than those of'Housui' at tetrad stage, but were more than those of 'Housui' at uninucleate microsporestage.
8. Lead nitrate was used to locate Ca~(2+)-ATPase in 'Niitaka' and 'Housui' anther. Theamount and distribution Ca~(2+)-ATPase changed during pollen development and weredifferent between 'Niitaka' and 'Housui'. Ca~(2+)-ATPase did not appear on microsporocytemembrane and a small amount Ca~(2+)-ATPase appeared some organelles membrane; theamount of Ca~(2+)-ATPase on cytoplasmic membrane and intine of microspore graduallyincreased during microspore development, but was less in cytoplasm. A large amount ofCa~(2+)-ATPase distributed on the cytoplasmic membrane, vacuole membrane and ubischbody of tapetal cell of 'Housui' from microeporocyte stage, the amount of Ca~(2+)-ATPasedecreased in disintegrating tapetum. The anthers were difference in Ca~(2+)-ATPasedistribution as compared with the same kind of cells at the similar developmental stage. TheCa~(2+)-ATPase did not appear on cytoplasmic membrane of microspore of 'Niitaka' at tetradstage. The amount of Ca~(2+)-ATPase on cytoplasmic membrane of microspore gradually increased, but was less than that of "Housui'; Ca~(2+)-ATPase was distributed on cytoplasmicmembrane and ubisch body of tapetum of 'Niitaka', and the amount of Ca~(2+)-AYPase wassimilar between 'Niitaka' and 'Housui' except microsporocyte stage.
9. Indole-3-acetic acid (IAA) content in floral bud of 'Housui' was very significantlyhigher than that of 'Niitaka' after tetrad stage. At the stage of pollen abortion, namely fromearly uninucleate pollen stage to late uninucleate pollen stage, IAA content of "Housui"was 3.70- 4.47 folds of 'Niitaka'. IAA deficiency of 'Niitaka' might be one of reason thatleads to pollen abortion; IAA oxidase activity in floral bud in prophase of microsporedevelopment of 'Niitaka' was significantly higher than that of 'Housui'. Change of PODactivity of both pear cultivars was very similar, and difference of POD activity was notsignificant, suggesting that POD was not key enzyme that leads to IAA deficiency.Abscisic acid (ABA) content of 'Niitaka' was significantly higher than that of'Housui' fromtetrad to binucleate stage.
10. The variational range of contents of both free Putrescine (Put) and Spermidine (Spd)in 'Niitaka' was small during floral bud development, whereas, variation of free Put andSpd content of 'Housui' showed a peak from early uninucleate pollen stage to lateuninucleate pollen stage. The free Put and Spd contents in Housui were higher than those of'Niitaka' at different developmental stage, especially at the stage of pollen abortion, thefree Put and Spd contents were 3-5 folds of 'Niitaka'. Free Spermine (Spm) content in'Niitaka' was significantly lower than that of 'Housui'. The contents of perchloric acidcovalently soluble conjugated (PSCC) Put, Spd and Spm of 'Housui' gradually increased,but the contents of PSCC Put, Spd and Spin in 'Niitaka' showed a peak that did not appearin 'Housui' during pollen abortion or ahead. The increase of PSCC polyamines in 'Niitaka'might result in the decrease of free polyamines and affect on pollen development.Variations of the content of perchloric acid covalently insoluble conjugated (PISCC)polyamine in two cultivars were similar. Total contents of different forms of Put, Spd andSpin did not have significant differences before pollen abortion, but were significant lowerin 'Niitaka' than that of 'Housui' after pollen abortion. The free Put, Spd and Spin were thepredominant forms in total contents in 'Housui', however, the predominant forms in Niitakawere PSCC Put, Spd and Spm.
In summary, the male sterile mechanism of 'Niitaka' was well characterized atmorphologic, ultra-structural, tyro-chemical, physiological and biochemical level. The present research clarified the stage of abortion of male sterile material and its maincytological mechanism, and further found out its abortive mechanism by way of physiologyand cyto-ehemistry. All these studies provided the basis for further research of molecularmechanism of'Niitaka' and the regulation of fertility expression.
1．‘新高’花药皱缩，发白，虽可以正常开裂，但花粉很少。在少数花药的横切面上虽然可看到微量花粉，但花粉染色很浅，且不规则，没有生活力。‘新高’花粉母细胞能正常进行减数分裂产生四分体；在小孢子产生后，发育缓慢，小孢子壁薄，加厚不明显；随着单核小孢子的液泡化，小孢子粘连降解，或小孢子细胞壁破损，细胞质、细胞核降解消失，很少见到二核花粉。说明‘新高’属于小孢子退化类型的雄性不育品种，小孢子退化主要发生在单核小孢子中期到晚期。
2．在花粉母细胞减数分裂前，‘新高’花粉母细胞和花药壁的发育与‘丰水’没差异，药壁包含表皮、药室内壁、中层和绒毡层，花粉母细胞被胼胝质包围。‘新高’四分小孢子染色很浅，胼胝质解体后，‘新高’较‘丰水’小孢子浅染，外粉壁发育迟缓而显得较薄。单核中期大部分小孢子收缩变形，且大小不等，细胞质解体，大部分不见细胞核。随后，小孢子壁破损，内含物外流，小孢子逐渐解体成为空壳或仅留下小孢子壁降解的残余物。在游离小孢子形成时，‘新高’绒毡层出现了不均匀的染色，其它花药壁结构与‘丰水’差异不明显。
3．‘新高’在小孢子母细胞时期出现包裹小孢子母细胞的胼胝质厚薄不均匀，部分不完整，胞内线粒体等细胞器的结构模糊，内质网分布少等早期异常现象；在四分体小孢子内小囊泡分布少，线粒体等细胞器的膜呈透明状，之后小孢子液泡膜破裂，细胞质开始解体，线粒体、质体、内质网等细胞器相继解体或退化，核仁解体，最后出现了同心环状的膜状体，以及细胞质及细胞核降解的残骸。而‘丰水’小孢子母细胞和雄配子体的线粒体结构清晰，液泡膜完整，且含有丰富的内质网、核糖体和高尔基体等细胞器，此外在四分体小孢子膜内分散着大量的小囊泡，这些小泡可能与原外壁的形成有关。
4．‘新高’四分小孢子表现出局部形成原外壁，或质膜和原外壁分离等不良现象，同时在胞内有很少的小囊泡。在单核小孢子时期，外壁上不能沉积孢粉素或只有表面沉积孢粉素，形成了透明状的外壁或无基足层的外壁，即使有的可以形成完整的外壁，但较正常孢子的外壁薄。而‘丰水’在四分小孢子时期形成了良好的原外壁，在雄配子体发育过程中，其外壁迅速发育。在花粉成熟时，花粉壁由薄的内壁及由基足层、基柱、覆盖层共同组成的外壁构成。两个品种的小孢子母细胞或四分体时期的胼胝质均能正常形成，并能及时降解，没有差异；‘丰水’小孢子外壁的自发荧光物质随雄配子体的发育而逐渐增多，在单核小孢子后期，小孢子外壁上除萌发的孔沟外均匀地沉积大量的自发荧光物质。‘新高’在单核小孢子早期小孢子外壁的荧光物质与‘丰水’差异不大，之后从局部到整个小孢子表面呈透明状，随之变形和相互粘连，有的小孢子表面荧光物质呈网状分布等异常表现。
5．‘新高’和‘丰水’小孢子母细胞时期的绒毡层细胞都含有大量的线粒体、内质网等细胞器；随着减数分裂的进行，两个品种的绒毡层细胞质收缩，细胞间和内切相面形成了许多空腔，细胞内高尔基体和分泌小泡增多，内质网膨胀形成槽库结构；四分体时期，‘丰水’绒毡层内分泌小囊泡和槽库结构的内质网进一步增多，并向药室分泌大量的乌氏体，而‘新高’绒毡层细胞内的部分细胞器消失，内质网垛叠，线粒体等细胞器边缘呈透明状，并产生了大量的小液泡；在小孢子发育的过程中，‘新高’并没有像‘丰水’一样出现结构清晰的内质网、线粒体等细胞器，而且出现了一些形态各异、结构模糊的细胞器；在绒毡层细胞解体时，‘丰水’绒毡层细胞内主要为油体等结构，并呈极性分布。而‘新高’细胞内还含有槽库结构的内质网，并和其它结构一起呈无序分布。‘新高’和‘丰水’绒毡层中自发荧光物质都产生于小孢子母细胞时期，以后荧光增强。在四分体小孢子即将释放单核小孢子时，‘丰水’绒毡层细胞表面分布大量的自发荧光物质颗粒，而‘新高’表面的自发荧光物质颗粒较少。
6．‘新高’结果枝和结果母枝中Fe、Zn、Cu、Ca、Mg和K含量较‘丰水’低，尤其是结果枝中ca和zn的含量差异更大，只有Mn的差异不显著。从四分体时期开始到单核小孢子后期，‘新高’和‘丰水’梨在花蕾中各矿质营养含量总体上呈逐渐下降的趋势，但‘新高’中Fe、zn、cu、Mg和Ca含量均低于‘丰水’，差异达到显著水平，Mn含量在单核花粉期后显著地高于‘丰水’，K的差异不明显。
7．运用焦锑酸钾沉淀法，对‘新高’和‘丰水’花药发育过程中Ca~(2+)进行了细胞化学定位研究。‘新高’和‘丰水’小孢子母细胞膜表面均有Ca~(2+)淀颗粒的分布。在四分体时期，‘丰水’小孢子表面积累了大量的Ca~(2+)沉淀颗粒，而在‘新高’中基本没有Ca~(2+)沉淀颗粒。随着胼胝质的解体，在‘丰水’小孢子的液泡、外壁以及细胞膜等部位均积累了Ca~(2+)沉淀颗粒，‘新高’小孢子中的Ca~(2+)沉淀颗粒则更多地分布在细胞质以及一些细胞器上。‘新高’小孢子中的Ca~(2+)常分布与其花粉败育有关。从小孢子母细胞开始，‘丰水’花药绒毡层的径切向面和内切向面均分布着大小不等的Ca~(2+)沉淀颗粒。随着小孢子的产生和发育，Ca~(2+)沉淀颗粒分布的量逐渐增多。在单核晚期，Ca~(2+)沉淀颗粒又减少。但‘新高’在四分体时期绒毡层中Ca~(2+)沉淀颗粒明显比‘丰水’要少，而单核小孢子时期Ca~(2+)分布较‘丰水’多。
8．应用铅沉淀法，分析了花药发育过程中Ca~(2+)-ATPase的分布。‘丰水’小孢子母细胞膜上基本上没有Ca~(2+)-ATPase的分布，只是在一些细胞器上有少量的分布；单核小孢子发育过程中，‘丰水’小孢子质膜及内壁上的Ca~(2+)-ATPaS逐渐增加，但在胞质内分布较少。从小孢子母细胞时期开始，在‘丰水’绒毡层细胞膜和液泡膜等细胞器以及乌氏体上分布有大量的Ca~(2+)-ATPase，随绒毡层的解体而逐渐减少。自四分小孢子开始，‘新高’和‘丰水’花药Ca~(2+)-ATPase就有差别：‘新高’四分小孢子细胞膜几乎没有Ca~(2+)-ATPase的分布，随小孢子的发育质膜上Ca~(2+)-ATPase虽逐渐增多，但比‘丰水’明显少；‘新高’绒毡层膜以及乌氏体有明显Ca~(2+)-ATPase分布，除小孢子母细胞时期外，与‘丰水’的差异不明显。
9．‘新高’梨花蕾中IAA含量从四分体后极显著地低于‘丰水’，在单核花粉早期和单核花粉晚期仅为‘丰水’的22．7％和27．0％，‘新高’梨IAA含量的亏缺可能是导致‘新高’败育的原因之一；‘新高’梨在花粉发育的前期IAA氧化酶活性显著地高于‘丰水’，而两个品种POD活性变化一致，并且活性的差异不显著，说明POD不是导致‘新高’IAA含量的亏缺的关键酶；‘新高’梨ABA含量从四分体以后却显著地高于‘丰水’。
10．‘新高’花蕾发育过程中游离态的Put和Spd变化幅度不大，‘丰水’则在单核花粉到二核花粉时期出现高峰，其含量是‘新高’的3-5倍；‘新高’中游离态的Spm也显著地低于‘丰水’。‘丰水’中高氯酸可溶共价结合态Put、Spd和Spm含量逐渐增加，‘新高’在花粉败育期或败育前期出现不同于‘丰水’的高峰，此种形态多胺含量的增加可能影响到游离态多胺含量的增加，而影响到花粉的发育。‘新高’和‘丰水’中高氯酸不可溶结合态多胺含量的变化基本一致。‘丰水’中不同种类的多胺均以游离态为主，‘新高’以高氯酸可溶结合态为主。
综上所述，本研究从细胞形态、亚细胞水平、细胞化学和生理生化水平上首次对‘新高’雄性不育机理进行了系统的研究。明确了‘新高’花粉败育的时期和主要细胞学机制，探明了‘新高’雄性不育在细胞化学和某些生理生化方面的败育机理。为从分子水平来研究‘新高’雄性不育的机理和人工调控育性的表达提供了理论依据。
Pear (Pyrus) belongs to gametophytic self-incompatibility, and allogamy among somevarieties also occur incompatibility. So, the suitable pollination trees and artificalsupplementary pollination are employed to meet expectant yield and quality. Some malesterile varieties of pear in production, for example 'Niitaka', 'Atagi', 'whangkeumbae', andso on, increase difficulties in choice of pollination tree and management. In order toconquer negative influence of male sterile cultivars, utilize potential cast of male sterileresource to solve pollen xenia effect and improve fruit quality, male sterile mechanism ofpear was investigated with male sterile 'Niitake' and male fertile 'Housui' (Pyrus pyrifolaiNakai). In the present study, transmission electron microscope, fluorescent and opticalmicroscope and some modem analytic equipment were employed, and the technology offluorescence labeling and cyto-chemistry was introduced. Based on identification of malesterility of 'Niitaka', pollen abortive stage and cytological characteristics, and distributionof fluorescent material were studied during pollen abortion. Moreover, Ca~(2+) andCa~(2+)-ATPase in anther were located to indentify the relationship between their distributionand pollen abortion. In addtion, hormones and polyamines in floral bud were analyzed tofurther study physiological mechanism of male sterility in pear. Main results of presentstudy are as follows:
1. Anther of 'Niitaka' appeared shrinkage and whitening, and could normally take placedehiscence, but had a small quantity of pollen. There was a lack of pollen on transversesection of anther, or a few pollens were found on a few sections, but these pollens appearedlight dyeing, abnormal shape and lower germination rate. The microspore mother cell couldnormally occur meiosis and produce tetrad; Development of microspore was slow, and wallof microspore was thin and slowly thicken after microsporogenesis; Microspore adhered toeach other, or wall, cytoplasm and nucleus of microspore was disintegrated with microspore vacuolization, and two-called pollen was not observed. It was suggested that 'Niitaka'belong to male sterile cultivar of microspore degenerated form, pollen abortion of 'Niitaka'appeared from middle unimicrospore stage to late unimicrospore stage.
2. Technology of paraffin section was used to observe anther development. There was notdifference of microspore mother and anther wall of 'Niitaka' and 'Housui' before meiosis,and anther wall were composed of epidermis, endothecium, middle layer and tapetum,microspore mother cell was surrounded by callose wall. Dyeing of microspore of 'Niitaka'was very light in tetrad, and was still light after callose wall disintegrated. Wall ofmicrospore slowly thickened and was very thin. Most microspores shnmk, deformed andappeared different size, cytoplasm was disintegrated, and nucleus disappeared. Microsporebecame hollow shell or debris with wall damaged and inclusion spilled. Tapetal cellappeared irregular dyeing at the stage of microspore just separated from tetrad, and othercomposition of anther wall disappeared difference between 'Niitaka' and 'Housui'.
3. Microsporocyte and male gametophyte of 'Housui' had perfect mitochondria,integrated vacuole membrane and rich endoplasmic reticulum, ribosome and Golgi bodyunder transmission electron microscope. There were abundant vesicles in microspore attetrad. Callose wall which surrounded microsporocyte of 'Niitake' appeared irregulardistribution, microsporocyte showed abnormal appearance, for example indiscerniblemitochondria structure, a small quantity of endoplasmic reticulum and so on; There were afew vesicle and some organdies of transparent membrane like mitochondria in microsporeof tetrad. Cytoplasm begun to disintegrate with vacuole membrane damaged, followingmitochondria, plastid and endoplasmic reticulum disintegrated, and nucleolus wasdegenerated. At last, there were some membrane bodies of concentric ring and debris ofcytoplasm and nucleus degenerated in microspore.
4. Microspore of 'Housui' could form perfect primexine at the tetrad stage, and couldrapid produce exine with male gametophyte development. Wall of mature pollen werecomposed of thin intine and exine including foot layer, baculum and tectum. Microspore of'Niitaka' appeared a portion of primexine or primexine separated from membrane at thetetrad stage, and there were a small quantity of vesicle. Microspore showed lacksporopollenin in exine or a little sporopollenin on the surface of exine, so exine oftransparency or lack foot layer was formed. Although some microspores could formintegrated exine, exine was thinner than that of normal microspore. Two cultivars disappeared difference in callose wall of formation and disintegration, which surroundedmicrosporocyte and microspore at tetrad stage. Auto-fluorescent material of microsporewall gradually increased with male gametophyte development in 'Housui', andauto-fluorescent material homogeneously distributed on wall of microspore except forcolporate at late microspore stage. There was not obvious difference in auto-fluorescentmaterial on wall of microspore between 'Housui' and 'Niitaka' at early microspore stage,microspore surface appeared transparency from portion to whole with auto-fluorescentmaterial decrease, and microspore deformed and adhered to each other, auto-fluorescentmaterial showed reticular distribution on the surface of some microspore which had beendisintegrated.
5. Tapetal cell of 'Niitaka' and 'Housui' had abundant mitochondria and endoplasmicreticulum at microsporocyte stage; cytoplasm of tapetal cell shrunk with rnicrosporocytemeiosis of 'Niitaka' and 'Housui', lots of cavity was formed in intercellular space and innersurface of tapetum with tapetal cell shrinkage. Golgi body and vesicle gradually increasedin tapetal cell, endoplasmic reticulum became dilated cistamae. The number of excretivevesicle and endoplasmic reticulum with cistarae further increased in tapetal cell of 'Housui',and tapetal cell excreted lots of ubish body for locule at tetrad stage. There was someabnormal appearance in tapetal cell of 'Niitaka' at tetrad stage, including part organellesdegenerated, stacked endoplasmic reticulum, mitochondria of transparent edge and a lot ofvacuoles. Tapetal cell of 'Niitaka' did not have perfect mitochondria and endoplasmicreticulum as tapetal cell of 'Housui' had, and had deformed and indiscernible organelleswith microspore vacuolization; lipid body appeared polar distribution in tapetal cell of'Housui' when tapetal cell disintegrated. Endoplasmic reticulum with cistarae and othermaterial appeared irregular distribution in tapetal cell of 'Niitaka'. Auto-fluorescentmaterial of tapetum appeared at microsporocyte, and fluorescent density strengthened withtapetal development in two cultivars. A mass of auto-fluorescent material which wasaccumulated on the surface of tapetal cell of 'Housui' was released and regularly distributedon the surface of microspore. But auto-fluorescent material of 'Niitaka' was less than thatof 'Housui', irregularly distributed on the surface of microspore.
6. The contents of iron (Fe), zinc (Zn), copper (Cu), calcium (Ca), magnesium (Mg) andpotassium (K) in basal bearing shoot and bearing shoot of 'Niitaka' were lower than thoseof 'Housui', especially the differences of the content of Ca and Zn in bearing shoot were even more between two cultivars, but the differences of the manganese (Mn) content wasnot significant. Mineral nutrition content in floral bud of 'Niitaka' and 'Housui' appeareddescendant trend from tetrad stage to late uninucleate microspore stage, and the contents ofFe、Zn、Cu、Mg and Ca of 'Niitaka' was significantly lower than those of'Housui', the Mncontent was significantly higher than those of 'Housui, the difference of K content was notsignificant.
7. Potassium antimontate was used to locate Ca~(2+) in 'Niitaka' and 'Housui' anther. Theamount and distribution of calcium precipitates changed during pollen development andwere different between 'Niitaka' and 'Housui'. The calcium precipitates accumulated on thesurface of microsporocyte of 'Niitaka' and 'Housui'. Lots of calcium precipitates appearedon the surface of microspore of 'Housui' at tetrad stage, but did not appeared on the surfaceof microspore of 'Niitaka'. The calcium precipitates gradually accumulated on thecytoplasmic membrane, vacuole membrane and exine of microspore of 'Housui' aftercallose wall disintegrated, but the most calcium precipitates were distributed in cytoplasmand some organelles of microspore of 'Niitaka'. The calcium precipitates appeared on thesurface of tapetal cell of 'Housui' at microsporocyte stage, and gradually increased withmicrosporogenesis and development, decreased at late uninucleate microspore stage. Thecalcium precipitates on the surface of tapetal cell of 'Niitaka' were less than those of'Housui' at tetrad stage, but were more than those of 'Housui' at uninucleate microsporestage.
8. Lead nitrate was used to locate Ca~(2+)-ATPase in 'Niitaka' and 'Housui' anther. Theamount and distribution Ca~(2+)-ATPase changed during pollen development and weredifferent between 'Niitaka' and 'Housui'. Ca~(2+)-ATPase did not appear on microsporocytemembrane and a small amount Ca~(2+)-ATPase appeared some organelles membrane; theamount of Ca~(2+)-ATPase on cytoplasmic membrane and intine of microspore graduallyincreased during microspore development, but was less in cytoplasm. A large amount ofCa~(2+)-ATPase distributed on the cytoplasmic membrane, vacuole membrane and ubischbody of tapetal cell of 'Housui' from microeporocyte stage, the amount of Ca~(2+)-ATPasedecreased in disintegrating tapetum. The anthers were difference in Ca~(2+)-ATPasedistribution as compared with the same kind of cells at the similar developmental stage. TheCa~(2+)-ATPase did not appear on cytoplasmic membrane of microspore of 'Niitaka' at tetradstage. The amount of Ca~(2+)-ATPase on cytoplasmic membrane of microspore gradually increased, but was less than that of "Housui'; Ca~(2+)-ATPase was distributed on cytoplasmicmembrane and ubisch body of tapetum of 'Niitaka', and the amount of Ca~(2+)-AYPase wassimilar between 'Niitaka' and 'Housui' except microsporocyte stage.
9. Indole-3-acetic acid (IAA) content in floral bud of 'Housui' was very significantlyhigher than that of 'Niitaka' after tetrad stage. At the stage of pollen abortion, namely fromearly uninucleate pollen stage to late uninucleate pollen stage, IAA content of "Housui"was 3.70- 4.47 folds of 'Niitaka'. IAA deficiency of 'Niitaka' might be one of reason thatleads to pollen abortion; IAA oxidase activity in floral bud in prophase of microsporedevelopment of 'Niitaka' was significantly higher than that of 'Housui'. Change of PODactivity of both pear cultivars was very similar, and difference of POD activity was notsignificant, suggesting that POD was not key enzyme that leads to IAA deficiency.Abscisic acid (ABA) content of 'Niitaka' was significantly higher than that of'Housui' fromtetrad to binucleate stage.
10. The variational range of contents of both free Putrescine (Put) and Spermidine (Spd)in 'Niitaka' was small during floral bud development, whereas, variation of free Put andSpd content of 'Housui' showed a peak from early uninucleate pollen stage to lateuninucleate pollen stage. The free Put and Spd contents in Housui were higher than those of'Niitaka' at different developmental stage, especially at the stage of pollen abortion, thefree Put and Spd contents were 3-5 folds of 'Niitaka'. Free Spermine (Spm) content in'Niitaka' was significantly lower than that of 'Housui'. The contents of perchloric acidcovalently soluble conjugated (PSCC) Put, Spd and Spm of 'Housui' gradually increased,but the contents of PSCC Put, Spd and Spin in 'Niitaka' showed a peak that did not appearin 'Housui' during pollen abortion or ahead. The increase of PSCC polyamines in 'Niitaka'might result in the decrease of free polyamines and affect on pollen development.Variations of the content of perchloric acid covalently insoluble conjugated (PISCC)polyamine in two cultivars were similar. Total contents of different forms of Put, Spd andSpin did not have significant differences before pollen abortion, but were significant lowerin 'Niitaka' than that of 'Housui' after pollen abortion. The free Put, Spd and Spin were thepredominant forms in total contents in 'Housui', however, the predominant forms in Niitakawere PSCC Put, Spd and Spm.
In summary, the male sterile mechanism of 'Niitaka' was well characterized atmorphologic, ultra-structural, tyro-chemical, physiological and biochemical level. The present research clarified the stage of abortion of male sterile material and its maincytological mechanism, and further found out its abortive mechanism by way of physiologyand cyto-ehemistry. All these studies provided the basis for further research of molecularmechanism of'Niitaka' and the regulation of fertility expression.
引文
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