平基槭花性别分化的细胞形态学观察

平基槭为杂性花,雄花与两性花同株,本文对其花性别分化过程进行了细胞形态学观察。结果发现,在花性别分化的早期,雄花和两性花的花芽中雌、雄蕊原基均具备,只是在花芽发育到一定时期,雄花的雌蕊原基发生选择性败育,败育发生在大孢子母细胞减数分裂为4个大孢子时期。两性花的雌蕊可以正常膨大结实,雄蕊花药虽然可以形成二核花粉,但不能正常开裂,属于不育雄蕊。初步分析认为,两性花雄蕊花药不能正常开裂与花粉囊壁纤维层木质化程度低有关。     

不同品种杏的性器官发育和结实性比较研究

通过对原产于不同地区的５１个杏品种的性器官发育和结实性的系统调查和测定,结果表明,１．花粉上和形状少数虽有一定差异,而大数色大多数品种相似,吕种间花粉萌发率尽管差异很大,但所有品种均为花粉可育型。２．大多数品种不完全花百分率很高,自然授粉率很,其主要原因是雌蕊败育。３．品种间不完全花百分率与花粉萌发率呈正相关;４．欧洲生态群品种与中国生态群品种比较芬粉大小和形态基本相似,而花粉萌发率和不安全花百分     

2个果梅品种雌蕊分化进程及相关生化指标分析

为了解果梅(Prunus mume Sieb.et Zucc.)雌蕊分化进程及其败育机制,采用石蜡切片法观察了不同时期果梅品种‘龙眼’(‘Longyan’)和‘大嵌蒂’(‘Daqiandi’)花芽纵切面的解剖结构,并对2个品种不同时期花器官发育状况、花芽百分率、花芽纵径和横径以及花芽中的可溶性糖、可溶性蛋白质和淀粉含量进行了测定分析.结果显示:雌蕊分化期、雌蕊分化末期及盛花期,品种‘龙眼’的不完全花比例均显著小于品种‘大嵌蒂’,其中,盛花期‘龙眼’不完全花比例仅为5.0％,而‘大嵌蒂’不完全花比例高达76.3％.品种‘龙眼’雌蕊分化过程经历未分化期、分化初期、分化期及分化末期4个阶段,且最终有95.0％的花芽在分化末期能顺利形成完全花;品种‘大嵌蒂’雌蕊分化过程则包含未分化期、分化初期、分化期、解体期、解体后修复期和分化末期6个阶段,且仅有23.7％的花芽能形成完全花.雌蕊分化的不同阶段2个品种花芽纵径和横径的变化与其分化进程基本一致.品种‘龙眼’完全花的可溶性糖和可溶性蛋白质含量均高于品种‘大嵌蒂’的完全花和不完全花、淀粉含量则低于后两者;品种‘大嵌蒂’不完全花的可溶性糖和可溶性蛋白质含量最低、淀粉含量则最高,与2个品种的完全花有显著差异.综合分析结果表明:品种‘大嵌蒂’的花芽在12月中上旬停止伸长生长、雌蕊分化停滞直至逐渐解体,这一时期即为品种‘大嵌蒂’雌蕊败育的关键时期;导致果梅雌蕊选择性败育的原因可能与花芽中大分子营养物质的分解代谢有关.     

太行花性器官发育的研究——两性花中雌雄性器官发育对温度的不同要求

用太行花（Ｔａｉｈａｎｇｉａ ｒｕｐｅｓｔｒｉｓ Ｙ汃ｅｔＬｉ）试管花进行雌雄蕊发育的温度试验表明：１． 雄蕊和雌蕊的发育对温度的要求是不同的,雄蕊发育的最适温度范围偏低,接近于１～６ ℃,高于１８ ℃雄蕊全部败育。雌蕊发育的最适温度范围偏高,约６～２６ ℃,低于３ ℃,大部分雌蕊在分化早期停止发育。２． 由高温引起的雄蕊败育过程始于雄蕊原基时期,这个过程一旦启动,雄蕊败育将不会因重新将它转入低温而逆转。通过控制实验温度获得了单性雄花（１～３ ℃）、单性雌花（６～２６ ℃）和两性花（６～１２ ℃）。实验结果将为性别控制和雄性不育研究开拓新思路     

山鸡椒雄花花芽发育形态解剖特征观察

采用体视显微镜、扫描电镜和石蜡切片技术对山鸡椒（Litsea cubeba（Lour.） Pers.）雄花花芽分化发育的外部形态和内部解剖结构进行了观察研究。结果显示:（1）山鸡椒雄花花芽分化发生可分为5个时期,即未分化期、花序原基分化期、苞片原基分化期、花原基分化期和花器官分化期,其中花器官分化期又可细分为花被原基分化期、雄蕊原基分化期和雌蕊原基分化期;各相邻分化时期存在一定重叠现象;花期从翌年1月上旬至3月下旬。（2）雄花成熟结构中具有独特的雄蕊蜜腺,蜜腺绿色且形态不规则,着生于内轮雄蕊基部,分布于花丝两侧,夹在内外轮雄蕊的花丝之间,与内轮花丝紧密相连。（3）雄蕊花药四室,花药壁发育属于基本型;腺质绒毡层;小孢子母细胞减数分裂过程中胞质分裂属于连续型;成熟花粉为2-细胞花粉粒;成熟花粉粒外壁刺突较多,刺突基部膨大,外壁露出部分粗糙,无薄壁区,有少数小穿孔。（4）山鸡椒雄花中绝大多数雌蕊发育至腹缝线卷合形成子房室时停止,柱头发育不良或者败育,花柱缩短或缺失,不能受精,直到开花结束,即发生退化。本研究明确了山鸡椒雄花花芽发育发生各个阶段时间、形态变化特点及外部形态变化特征,山鸡椒小孢子发生、雄配子体发育至散粉期变化特点和规律以及雄花中退化雌蕊发育的进程,可为山鸡椒优良品种选育、调控花期和提高结实率提供一定的参考。     

丹桂和籽银桂花芽分化的研究

采用石蜡切片技术, 对不育的丹桂(Osmanthus fragrans ‘Dangui’ )和可育的籽银桂(Osmanthus fragrans ‘Ziyingui’)花芽分化的过程进行了研究。结果表明, 桂花(Osmanthus fragrans Lour.)花芽形态分化可分为花芽分化初始期、总苞分化期、花原基分化期、顶花花被分化期、雄蕊分化期和雌蕊分化期6个阶段。雄蕊的发育在丹桂和籽银桂之间基本没有区别, 都能形成完整的花粉囊和成熟的花粉粒。但是, 雌蕊的发育在可育的籽银桂与不育的丹桂之间存在明显差异。根据花芽分化的过程证明, 丹桂的不育是由于雌蕊发育不正常导致的。     

丹桂和籽银桂花芽分化的研究

采用石蜡切片技术,对不育的丹桂（Osmanthus fragrans ‘Dangui’）和可育的籽银桂（Osmanthus fragrans ‘Ziyingui’）花芽分化的过程进行了研究。结果表明,桂花（Osmanthus fragrans Lour.）花芽形态分化可分为花芽分化初始期、总苞分化期、花原基分化期、顶花花被分化期、雄蕊分化期和雌蕊分化期6个阶段。雄蕊的发育在丹桂和籽银桂之间基本没有区别,都能形成完整的花粉囊和成熟的花粉粒。但是,雌蕊的发育在可育的籽银桂与不育的丹桂之间存在明显差异。根据花芽分化的过程证明,丹桂的不育是由于雌蕊发育不正常导致的。     

濒危植物长柄双花木自然种群结实的花粉和资源限制

长柄双花木（Disanthus cercidifolius Maxim．vsr．longipes H．T．Chang）只零星分布于湖南、江西和浙江等省的少数县,已处于濒危状态。通过对其野外自然种群试验,研究了花粉和资源的有效性对雌性生殖的影响,并进一步探讨了长柄双花木“花多果少”的生殖机制。结果表明：不同来源的花粉对长柄双花木结果率及结籽率均产生显著影响。花粉来源而不是花粉数量对长柄双花木结实存在显著影响。自然条件下,长柄双花木“花一果转化率”约为54馏,“胚珠-种子转化率”约为83．19。施肥处理有利于花芽发育,可使植株花芽败育率明显下降,开花比例升高;同时座果率和结籽率均显著提高,而果实败育率则显著降低。随剪除叶片量的增加,结果率明显降低,但果实败育率在处理间无显著差异;剪除弱小枝条及病虫枝后,个体总开花数虽有所下降,但是结果率和结籽率均显著提高。去除花后,长柄双花木平均单果重、单粒种子重均显著降低。营养水平的时空异质性使自然种群结籽率受到资源和花粉的限制;补充资源可以通过增强花对传粉者的吸引而间接影响传粉。长柄双花木自然种群的坐果率极低,而且存在严重的“大小年”结果现象,不同的因子可能对其低水平坐果率产生相互作用。选择性败育、子房供应和雄性功能假说似乎是长柄双花木“花多果少”的生殖策略的最合理解释。     

罗汉果花芽分化过程中形态及其激素水平变化特征

采用石蜡切片和酶联免疫法(ELISA)对罗汉果雄性、雌性、两性花芽分化过程的形态和激素水平变化进行观测,为罗汉果开花调控和品种选育提供科学依据。结果表明:(1)罗汉果雄性、雌性、两性花的花芽分化过程均可分为花芽未分化期、花芽分化初期、花序分化期、萼片原基分化期、花瓣原基分化期、雄蕊原基分化期和雌蕊原基分化期7个阶段。雄蕊原基分化期前,3种花芽分化过程无明显差异,各时期形态特征均依次为:茎端呈圆锥状(花芽未分化期)→茎端经半球形变成扁平状(花芽分化初期)→距茎端5~7节位处分化出穗状花序(花序分化期)→小花原基周围形成5个萼片原基(萼片原基分化期)→萼片原基内侧形成5个花瓣原基(花瓣原基分化期)。雄蕊和雌蕊原基分化期,3种花芽分化过程存在明显差异,雄蕊原基内侧出现雌蕊原基后,雄花芽雄蕊原基继续发育成雄蕊,雌蕊原基停滞生长,退为一个小突起;雌花芽雌蕊原基继续发育成雌蕊,雄蕊原基生长缓慢,退化为小花丝;两性花芽雌蕊和雄蕊原基均继续发育,形成外观正常的雌蕊和雄蕊。(2)内源激素脱落酸(ABA)、赤霉素(GAs)和玉米素核苷(ZR)含量在3种花芽分化过程中变化规律相似,即ABA含量在花芽生理分化期降低,花芽形态分化期升高,而GAs和ZR含量则基本保持不变;吲哚乙酸(IAA)含量在3种花芽分化过程中变化存在明显差异,雌花芽IAA含量在花芽生理分化期升高,花芽形态分化期逐渐降低,而雄性和两性花芽的IAA含量则基本保持不变。ABA/GAs、ABA/IAA、ZR/IAA和ZR/GAs激素含量比值在3种花芽分化过程中变化规律相似,ABA/GAs在花芽生理分化期降低,花芽形态分化期升高,而BA/IAA、ZR/IAA和ZR/GAs则基本保持不变。研究认为,罗汉果花芽分化过程经历一个"两性期",高ABA含量和ABA/GAs比值有利于罗汉果花芽分化,IAA可能对罗汉果花性分化具有重要作用。     

温度对离体培养中风信子再生雄蕊的诱导,小孢子发生和花粉发育的影响

风信子(Hyacinthus orientalis)雄蕊再生的最适温度是25℃。小孢子发生和花粉发育是在温度不断下降的过程中完成的。母细胞分化的最适温度是20—25℃;减数分裂的最适温度是20℃;花粉第一次有丝分裂的最适温度是10℃。按这样的温度条件能够将再生雄蕊培养成熟,并且成熟雄蕊的花粉具有较高的萌发率。不合适的温度将导致小孢子发生和花粉发育停止在某个阶段,最终花粉败育。     

Pollen and resource limitations to lifetime seed production in a wild population of the endangered plant Disanthus cercidifolius var. longipes H. T. Chang (Hamamelidaceae)

Disanthus cercidifolius Maxim. var. longipes H. T. Chang, a plant species that only occurs in a few counties in Hunan, Jiangxi and Zhejiang Provinces and with a relatively small number of individuals, is ranked as a second Class endangered species for conservation in China. We have studied the effect of pollen and resources available to female reproduction, and the reproductive mechanism of “excess flowers with low fruit set” in Disanthus cercidifolius Maxim. var. longipes H. T. Chang was discussed. Results are as follows Pollen from different sources has significant effects on fruit set and seed set of Disanthus cercidifolius Maxim. var. longipes H. T. Chang. The pollen source rather than pollen numbers significantly affected reproduction of this species. In wild populations, producing one fruit needs about 54.8 flowers, and one satiation seed needs about 6.60 flowers or 83.19 ovules. After fertilizing, which was propitious to flower development, the abortion rate of flower buds was decreasing, but the flowering rate was increasing. The fruit set and seed set was also significantly increasing, while abortion rate of fruit was significantly decreasing. With the increasing percentages of cutting leaves, the fruit set decreased, but the abortion rate of fruit shows no significant differentiation among treatments. After cutting branches that were puny, broken and insectin-fested branches, the flower number seemed to be decreasing, but the fruit set and seed set all increased significantly. After removing some flowers, the fruit set was calculated with respect to the number of flowers remaining after the treatment increased with increasing of percentages of flower removal, whereas fruit set calculated with respect to the initial number of flowers remained constant, and the mean weights of per fruit and per seed all decreased significantly. Sufficient spatial or temporal heterogeneities in nutrient levels might allow limitation of seed set by resources and pollen in a natural population, while supplying resources may indirectly affect pollination by increasing attraction of the flowers to pollinators. There were very low fruit and seed sets in natural populations of Disanthus cercidifolius Maxim. var. longipes H. T. Chang. Different factors may have interacted to effect a low fruit set. A joint adoption of the “selection abortion hypothesis”, “ovary reserve hypothesis” and “male function hypothesis” seems to be the most likely explanation for the reproductive strategy of “excess flowers with few fruit sets” in Disanthus cercidifolius Maxim. var. longipes H. T. Chang. __________ Translated from Acta Ecologica Sinica, 2006, 26(2): 496–502 [译自: 生态学报]     

Flower bud differentiation and development in fruiting and non-fruiting shoots in relation to fruit set in apricot (Prunus armeniaca L.)

Situations of high flower bud drop and low fruit set without apparent causes are common in fruit trees. The term flower quality has been coined to explain differences among flowers in their capacity to set fruit, but the causes underpinning these differences are largely unknown. This lack of knowledge is based on the fact that these differences are established a posteriori and there are no criteria to determine a priori what will make a flower to set a fruit or to drop. In this work, we profit from the empirical knowledge that there are fruiting and non-fruiting shoots to explore to which extent flower bud differentiation and bud development will affect the subsequent fruit set. For this purpose, the processes from flower bud differentiation to fruit set were sequentially analyzed in both types of shoots, over 2 years. More than half of the buds from long shoots aborted development and dropped before flowering. At anthesis, most of the remaining flowers showed underdeveloped pistils that failed to sustain pollen germination or pollen tube growth along the pistil. This unsuccessful development resulted in clear differences in fruit set between both types of branches. These results highlight that flower bud differentiation and development play an important role for fruit set and that developmental timing appears critical to reach anthesis with a fully developed pistil.     

Pollen and resource limitations to lifetime seed production in a wild population of the endangered plant Disanthus cercidifollus var.Iongipes H.T.Chang (Hamamelidaceae)

Disanthus cercidifolius Maxim.var.longipes H.T.Chang,a plant species that only occurs in a few counties in Hunan,Jiangxi and Zhejiang Provinces and with a relatively small number of individuals,is ranked as a second Class endangered species for conservation in China.We have studied the effect of pollen and resources available to female reproduction,and the reproductive mechanism of "excess flowers with low fruit set" in Disanthus cercidifolius Maxim.var.longipes H.T.Chang was discussed.Results are as follows:Pollen from different sources has significant effects on fruit set and seed set of Disanthus cercidifolius Maxim.var.longipes H.T.Chang.The pollen source rather than pollen numbers significantly affected reproduction of this species.In wild populations,producing one fruit needs about 54.8 flowers,and one satiation seed needs about 6.60 flowers or 83.19 ovules.After fertilizing,which was propitious to flower development,the abortion rate of flower buds was decreasing,but the flowering rate was increasing.The fruit set and seed set was also significantly increasing,while abortion rate of fruit was significantly decreasing.With the increasing percentages of cutting leaves,the fruit set decreased,but the abortion rate of fruit shows no significant differentiation among treatments.After cutting branches that were puny,broken and insectinfested branches,the flower number seemed to be decreasing,but the fruit set and seed set all increased significantly.After removing some flowers,the fruit set was calculated with respect to the number of flowers remaining after the treatment increased with increasing of percentages of flower removal,whereas fruit set calculated with respect to the initial number of flowers remained constant,and the mean weights of per fi'uit and per seed all decreased significantly.Sufficient spatial or temporal heterogeneities in nutrient levels might allow limitation of seed set by resources and pollen in a natural population,while supplying resources may indirectly affect pollination by increasing attraction of the flowers to pollinators.There were very low fruit and seed sets in natural populations ofDisanthus cercidifolius Maxim.var.longipes H.T.Chang.Different factors may have interacted to effect a low fruit set.A joint adoption of the "selection abortion hypothesis","ovary reserve hypothesis" and "male function hypothesis" seems to be the most likely explanation for the reproductive strategy of"excess flowers with few fruit sets" in Disanthus cercidifolius Maxim.var.longipes H.T.Chang.     

不同生境与花部特征对巫山淫羊藿结实特性的影响

对不同生境中巫山淫羊藿结实特性进行比较研究。结果表明:(1)三个种群在繁殖时期中不同花部特征的植株繁殖投资差异不大,如每株分枝数、每株花序数和每株开花数差异均不显著;(2)三个种群中每株结实数、每株结实率、每花序结实数在不同花部特征中差异显著,PLS和SAA要显著大于PESS;(3)三种花部特征在不同生境中单株分枝数和单株花序数均无显著差异。在不同花部特征中种群1开花数量相较少,因而在结实方面均值均要小于种群2和种群3;(4)不同花部特征的果实饱满种子数、败育数和败育率均具有显著差异,而种子数差异不大,但在种群间均无显著的差异。说明种子的多少更多地受到花部特征的影响。总之,巫山淫羊藿的结实特性受到生境和花部特征的双重影响,其中花型比生境对巫山淫羊藿结实特性的影响更为明显。     

云南草寇花序内不同部位的性分配

两性花植物花序内的性分配常存在差异,资源竞争、结构效应、交配环境(雌雄异熟、传粉者定向访花行为等)或授粉不均匀等几种假说可以解释这种现象。为验证上述假说,该研究以云南草寇两种表型(雄先熟型和雌先熟型)为材料,分析了其花序内不同部位(基部、中部和顶部)的每花花粉数、胚珠数、花粉/胚珠比、结实率和结籽率,花序内传粉者的定向访花行为,以及人工辅助授粉和去花处理对结实率和结籽率的影响。结果表明:两种表型花序内每花花粉数不随部位而变化,每花胚珠数、结实率和结籽率由基部到顶部依次降低,每花花粉/胚珠比由基部到顶部依次增加,表明顶部花存在偏雄的性分配。人工辅助授粉后,结实率、结籽率仍由基部到顶部依次降低,表明授粉不均匀假说不能解释云南草寇花序内不同部位结实率、结籽率的差异。去除基部和中部花后,顶部花人工辅助授粉条件下的结实率、结籽率与基部花人工辅助授粉条件下的结实率、结籽率无差异,表明云南草寇花序内不同部位结实率、结籽率的差异主要由资源竞争引起。雌先熟表型每花花粉数、花粉/胚珠比高于雄先熟表型,表明两种表型存在性分配差异。传粉者主要先访问云南草寇基部的花,然后向顶部移动。云南草寇花序内顶部偏雄的性分配可能是由资源竞争和传粉者定向访花造成的。     

斗竹的开花生物学特性研究

通过野外观测及光学解剖,观察了斗竹(Oligostachyum spongiosum)开花林相、开花动态、花器官构造、结实情况,以及花后林相更新等生物学特性,采用光学显微技术结合石蜡制片,对斗竹的大、小孢子的发生及雌、雄配子体的发育过程进行研究。结果表明：(1)斗竹为一次性整体开花竹类,花期为4月下旬～5月下旬,花期约持续45 d,成花量大。(2)花序为圆锥状混合花序,每花序由4枚小穗构成;小穗细长,每枚小穗由5～17枚小花组成;小花为颖花,顶部小花不发育,外稃、内稃各1枚;浆片3枚,卵圆形;雄蕊4～6枚(多为6),每枚花药具有4个花粉囊,花药壁发育为基本型,绒毡层为腺质型,小孢子四分体为左右对称型,成熟花粉粒为2 细胞型,球形,表面纹饰颗粒状,具单个萌发孔,花粉发育过程中部分花药出现异常收缩及空腔的败育花粉粒;雌蕊1枚,柱头3叉,羽毛状,子房1室,胚珠倒生,厚珠心,胚囊为蓼型,成熟胚囊结构及发育过程均正常;雌雄同熟,异花授粉,果实为颖果。(3)斗竹花后全林死亡,结实率低,自然条件下结实率为8.1%。研究结果为研究竹子系统分类、开花机制,开展杂交育种及竹林更新复壮工作等提供基础性资料。     

内蒙古居群抗寒西伯利亚杏繁殖生态学研究

对内蒙古居群抗寒西伯利亚杏的12个种源同一树龄的102株进行雌蕊败育率、花粉量、花粉活力、柱头可授性及自交不亲和性进行研究,以明确西伯利亚杏主产地资源的花期生物学特性和繁殖特性,为杏杂交育种和资源利用提供理论依据。结果表明:(1)有86.27%的西伯利亚杏雌蕊败育率低于90%;73.53%的西伯利亚杏单朵花花粉量在1×104～5×104之间;86.27%的西伯利亚杏花粉活力低于50%,且寿命较短。(2)花苞在将开未开时,柱头已经具备一定的可授性,开花后1～4d内柱头可授性保持较高水平,第5天开始有下降趋势。(3)荧光显微观察结果显示,授粉后16h时,96.08%的杏花花粉管在花柱上部;授粉后24h时,74.51%的杏花花粉管仍在花柱上部;授粉后48h时,35.29%的杏花花粉管到达花柱中部;授粉后80h时,花粉管到达花柱下部的杏花约50%,但受精胚珠数为0。研究认为,西伯利亚杏基本为自交不亲和。     

绵枣儿花芽分化的形态解剖学研究

为探明绵枣儿(Barnardia japonica)在哈尔滨地区的年生长节律以及花芽分化进程,该文以从长白山引种至东北林业大学花卉研究所苗圃内的绵枣儿为材料,采用田间观察法研究绵枣儿的年生长节律,并采用石蜡切片法观察其花芽分化各阶段的形态解剖学特征。结果表明:(1)绵枣儿在哈尔滨地区的生长节律大致可以分为四个时期,即花芽分化与发育期、开花期、结实期、休眠期。(2)绵枣儿花芽分化进程可以分为七个阶段,即4月中上旬,由于土壤温度较低,鳞茎仍处于未分化期; 4月下旬进入花序原基分化期; 5月上旬苞片原基分化; 5月下旬为小花原基分化期; 5月末至6月初花被片原基分化; 6月上旬进入雄蕊原基分化期; 6月下旬为雌蕊原基分化期。该研究明确了绵枣儿在哈尔滨地区的年生长节律和花芽分化各阶段的解剖学特性,为园林应用和新品种的选育提供了一定的科学依据。     

Temperatures during flower bud development affect pollen germination,self‐incompatibility reaction and early fruit development of clementine (Citrus clementina Hort. ex Tan.)

• One of the key environmental factors affecting plant reproductive systems is temperature. Characterising such effects is especially relevant for some commercially important genera such as Citrus. In this genus, failure of fertilisation results in parthenocarpic fruit development and seedlessness, which is a much‐prized character. Here, we characterise the effects of temperature on flower and ovary development, and on pollen–pistil interactions in ‘Comune’ clementine (Citrus clementina Hort. ex Tan.).
• We examine flower bud development, in vitro pollen germination and pollen–pistil interaction at different temperatures (15, 20, 25 or 30 °C). These temperatures span the range from ‘cold’ to ‘hot’ weather during the flowering season in many citrus‐growing regions.
• Temperature had a strong effect on flower and ovary development, pollen germination, and pollen tube growth kinetics. In particular, parthenocarpic fruit development (indicated by juice vesicle growth) was initiated early if flowers were exposed to warmer temperatures during anthesis.
• Exposure to different temperatures during flower bud development also alters expression of the self‐incompatibility reaction. This affects the point in the pistil at which pollen tube growth is arrested and confirms the role of sub‐ and supra‐optimal temperatures in determining the numbers of pollen tubes reaching the ovary.