单性木兰（木兰科）花的形态发生

报道了单性木兰(Kmeria septentrionalis Dandy)花的形态发生过程。发现过去一直被认为是雌花条状披针形的“内轮花被片”,实际为退化雄蕊,它形态发生的时间与位置均与雄花的雄蕊相同,在成熟结构中仍可见药室残迹,说明单性木兰的雌性花是由两性花退化而来。通过与K. duperreana(Pierre) Dandy和Magnolia thailandica Noot. &; Chalermglin雌花的比较,发现它们雌花的形态相同,从而得知人们长期以来对此3种植物雌花的认识有误,原一直认为的“内轮花被片”实为退化雄蕊。     

Study of homeosis in the flower of Philodendron (araceae): a qualitative and quantitative approach

This study deals specifically with floral organogenesis and the development of the inflorescence of Philodendron squamiferum and P. pedatum. Pistillate flowers are initiated on the lower portion of the inflorescence and staminate flowers are initiated on the distal portion. An intermediate zone consisting of sterile male flowers and atypical bisexual flowers with fused or free carpels and staminodes is also present. This zone is located between the sterile male and female floral zones. In general, the portion of bisexual flowers facing the male zone forms staminodes, and the portion facing the female zone develops an incomplete gynoecium with few carpels. The incomplete separation of some staminodes from the gynoecial portion of the whorl shows that they belong to the same whorl as the carpels. There are two levels of aberrant floral structures in Philodendron: The first one is represented by the presence of atypical bisexual flowers, which are intermediates between typical female flowers and typical sterile male flowers. The second one is the presence of intermediate structures between typical carpels and typical staminodes on a single atypical bisexual flower. The atypical bisexual flowers of P. squamiferum and P. pedatum are believed to be a case of homeosis where carpels have been replaced by sterile stamens on the same whorl. A quantitative analysis indicates that in both species, on average, one staminode replaces one carpel.     

Homeosis in Araceae Flowers: The Case of Philodendron melinonii

The early stages of development of the inflorescence of Philodendronmelinonii were examined using scanning electron microscopy.Pistillate flowers are initiated on the lower portion of theinflorescence and staminate flowers are initiated on the distalportion. The male flowers have four to five stamens. The femaleflowers have a multilocular ovary consisting of four to sixlocules. A transition zone consisting of sterile male flowersand bisexual flowers with fused or free carpels and staminodesis also present on the inflorescences. This zone is locatedbetween the male and female flower zones. Generally, the portionof the bisexual flower facing the male zone forms stamens, andthe portion facing the female zone develops an incomplete gynoeciumwith few carpels. In P. melinonii, the incomplete separationof staminodes from the gynoecial portion of the whorl showsthat the staminodes and carpels belong to the same whorl. Thebisexual flowers of P. melinonii are believed to be a case ofhomeosis where carpels have been replaced by sterile stamenson the same whorl. At the level of the inflorescence, pistillateand staminate flowers are inserted on the same contact parastichiesalong the inflorescence; there is no discontinuity between thefemale zone, the bisexual zone, and the male zone. The presenceof bisexual flowers is believed to correspond to a morphogeneticgradient at the level of the inflorescence as a whole. Copyright2000 Annals of Botany Company Flower, development, gradient, inflorescence     

Pistillate and staminate flower development in dioecious Pistacia vera (Anacardiaceae)

The development of staminate and pistillate flowers in the dioecious tree species Pistacia vera L. (Anacardiaceae) was studied by scanning electron microscopy with the objective of determining organogenetic patterns and phenology of floral differentiation. Flower primordia are initiated similarly in trees of both sexes. Stamen and carpel primordia are initiated in both male and female flowers, and the phenology of organ initiation is essentially identical for flowers of both sexes. Vestigial stamen primordia arise at the flanks of pistillate flower apices at the same time functional stamens are initiated in the staminate flowers. Similarly, a vestigial carpel is initiated in staminate flowers at the same time the primary, functional carpel is initiated in pistillate flower primordia. Differences between the two sexes become apparent early in development as, in both cases, development of organs of the opposite sex becomes arrested at the primordial stage. Male flowers produce between four and six mature functional stamens and female flowers produce a gynoecium with one functional and two sterile carpels.     

细枝柃性别表达特性及资源分配

已有的资料将柃木属(Eurya)描述为严格的雌雄异株植物, 性别变异现象极为少见。目前仅在柃木(E. japonica)和钝叶柃(E. obtusifolia)等少数种类中报道过两性花的存在。近几年笔者发现细枝柃(E. loquaiana)存在性别变异现象, 性别变异株上具有不同性别类型的花。该文从单花和植株水平分析了细枝柃的性别表达特性, 并对不同类型花的花部构件生物量分配进行比较分析。结果表明, 细枝柃具有6种类型的花, 从单花水平上看, 细枝柃性别有雌性、雄性及两性3种类型; 细枝柃性别在植株水平上体现较为复杂, 有雌株, 雄株, 雌花和两性花同株, 雄花和两性花同株, 雌雄异花同株及雌花、雄花、两性花同株6种类型; 在细枝柃花部构件生物量分配中, 雄花(包括雄株花和变异株雄花)花部构件生物量分配中雄蕊生物量的分配低于雌花(包括雌株花和变异株雌花)中雌蕊生物量的分配; 两性花中, 雄蕊生物量分配低于雌蕊, 这是其优化资源分配的手段, 进而获取最大适合度收益。     

The role of growth regulators in the differentiation of flowers and inflorescences

Growth regulators participate in the differentiation of floral parts, determining the developmental path of the respective type of inflorescence. The effect depends on the expression of the peculiarities of floral part differentiation, the recognition of the character of endogenous substances in certain stages and the choice of the suitable regulator for application. In the primitive flower ofPapaver petals and stamens are formed from the peripheral meristem with a lower content of auxins and a higher level of gibberellic substances. The pistil arises later from central tissues with a higher level of auxins and inhibitory substances. The stamens are more sensitive to the higher level of auxin substances, and by a suitable application of GA₃ and BAP they can be transformed into petals; in this way double flower forms arise. In the differentiation of floral parts ofCampanula, Rosa andMelandrium similar regularities assert themselves in time successions, but in another spatial arrangement. Sex differentiation of diclinous flowers ofMelandrium is based on differences in heterochromosomes XY and XX. The rise of the zygomorphic flower ofVeronica is accompanied by a different distribution of endogenous substances which affect the development of petals, stamens and the pistil. The differentiation of flowers in the racemose inflorescence occurs in the acropetal succession, and lateral primordia inCampanula develop into actinomorphic regular flowers, whereas inDigitalis they are zygomorphic and only the terminal flower is peloric. In the initial phases the staminate tassel and the pistillate ear in maize are identical. Earlier differentiation of the terminal pistillate tassel is connected with a higher level of gibberellins and the later development of the lateral pistillate ear is accompanied by the increase in auxin-like substances and inhibitions. Similar correlations were found in the development of staminate catkins and the differentiation of pistillate flowers in terminal buds ofJuglans regia. By the application of auxin-like substances it is possible to achieve the transformation of primordia of the staminate tassel into the pistillate ear in maize or to regulate the number of staminate catkins and pistillate flowers on twigs of the walnut tree. In the capitulum of the sunflower differences arise between peripheral pistillate ray flowers and hermaphrodite tubular ones. By applying GA₃ and BAP the number of ray flowers is increased. If the normal course of inflorescence differentiation is affected with a suitable type of regulator, a range of floral abnormalities appears which permit to assess the intervention in different developmental stages and the reaction of the primordium to the applied type of regulator. Abnormalities also suggest some phylogenetic correlations.     

Evidence of trimonoecy in Phyllanthaceae: Phyllanthus acidus

Pollen from staminate flowers and pistillate flowers with ??staminodes?? of Phyllanthus acidus Skeels were analyzed under scanning electron microscopy, and tests of pollen viability and in?vitro germination were carried out to verify possible similarities between the three types of flowers. The results show that pistillate flowers with ??staminodes?? are bisexual, indicating the occurrence of trimonoecy in this species.     

Gender variation in Actinidia deliciosa,the kiwifruit

Summary Flower and fruit characters were measured in ten female, five male and five fruiting male selections of A. deliciosa var deliciosa (A. Chev) Liang and Ferguson. Flowers from female vines had functional pistils, which contained many ovules. Stamens appeared to be fully developed but produced only empty pollen grains. Flowers from male vines had functional stamens that produced high percentages of pollen grains with stainable cytoplasmic contents. Pistils did not contain ovules and were generally small with vestigial styles. Fruiting male vines had both staminate and bisexual flowers. Staminate flowers were similar to those found on strictly male vines. Bisexual flowers produced ovules and stainable pollen. Pistils were smaller than in pistillate flowers. Although the three flower sexes differed in style length, ovary dimensions and ovules per carpel, staminate and bisexual flowers were similar in number of flowers per inflorescence, stamen filament length, pollen stainability, inflorescence rachis length and carpel number, and differed from pistillate flowers in these characters. The three flower sexes had similar sepal and petal numbers. The fruit of fruiting males were considerably smaller than those of females. Low ovule number appears to be the major factor limiting fruit size in the fruiting males studied. Prospects for developing hermaphroditic kiwifruit cultivars through breeding are discussed.     

Floral anatomy of Paepalanthoideae (Eriocaulaceae, Poales) and their Nectariferous structures

BACKGROUND AND AIMS: Eriocaulaceae (Poales) is currently divided in two subfamilies: Eriocauloideae, which comprises two genera and Paepalanthoideae, with nine genera. The floral anatomy of Actinocephalus polyanthus, Leiothrix fluitans, Paepalanthus chlorocephalus, P. flaccidus and Rondonanthus roraimae was studied here. The flowers of these species of Paepalanthoideae are unisexual, and form capitulum-type inflorescences. Staminate and pistillate flowers are randomly distributed in the capitulum and develop centripetally. This work aims to establish a floral nomenclature for the Eriocaulaceae to provide more information about the taxonomy and phylogeny of the family. METHODS: Light microscopy, scanning electron microscopy and chemical tests were used to investigate the floral structures. KEY RESULTS: Staminate and pistillate flowers are trimerous (except in P. flaccidus, which presents dimerous flowers), and the perianth of all species is differentiated into sepals and petals. Staminate flowers present an androecium with scale-like staminodes (not in R. roraimae) and fertile stamens, and nectariferous pistillodes. Pistillate flowers present scale-like staminodes (except for R. roraimae, which presents elongated and vascularized staminodes), and a gynoecium with a hollow style, ramified in stigmatic and nectariferous portions. CONCLUSIONS: The scale-like staminodes present in the species of Paepalanthoideae indicate a probable reduction of the outer whorl of stamens present in species of Eriocauloideae. Among the Paepalanthoideae genera, Rondonanthus, which is probably basal, shows vascularized staminodes in their pistillate flowers. The occurrence of nectariferous pistillodes in staminate flowers and that of nectariferous portions of the style in pistillate flowers of Paepalanthoideae are emphasized as nectariferous structures in Eriocaulaceae.     

花叶芋(天南星科)的花器官发生

利用扫描电镜首次观察了天南星科花叶芋(Colocasia bicolor) 的花器官发生过程。花叶芋的肉穗花序由无花被的单性花构成, 雌花发生于花序基部, 雄花发生于花序上部, 中性花位于花序中间部位。雄花: 3 或4 个初生雄蕊原基轮状发生, 随后每个初生原基一分为二, 形成6或8个次生原基; 一部分次生原基在其后的发育过程中融合, 形成5 或7 枚雄蕊; 雄花发育过程中未见雌性结构的分化; 花药的分化先于花丝; 雄蕊合生成雄蕊柱。雌花: 合生心皮, 3或4个心皮原基轮状发生, 未见雄性结构的分化。中性花来源于雌雄花序过渡带上, 属于雄蕊原基的滞后发育以及发育成熟过程中的退化; 与彩叶芋属(Caladium)不同, 此过渡区未见畸形两性花。初生雄蕊原基二裂产生次生原基的次生现象在目前天南星科花器官发生中显得比较特殊, 同时初步探讨了次生原基的融合方式。     

臭椿花器官分化的初步研究

利用扫描电镜(SEM)和光镜(LM)对臭椿花序及花器官的分化和发育进行了初步研究,表明:1)臭椿花器官分化于当年的4月初,为圆锥花序;2)分化顺序为花萼原基、花冠原基、雄蕊原基和雌蕊原基。5个萼片原基的发生不同步,并且呈螺旋状发生;5个花瓣原基几乎同步发生且其生长要比雄蕊原基缓慢;雄蕊10枚,两轮排列,每轮5个原基的分化基本是同步的;雌蕊5,其分化速度较快;3)在两性花植株中,5个心皮顶端粘合形成柱头和花柱,而在雄株中,5个心皮退化,只有雄蕊原基分化出花药和花丝。本研究着重观察了臭椿中雄花及两性花发育的过程中两性花向单性花的转变。结果表明,臭椿两性花及单性花的形成在花器官的各原基上是一致的(尽管时间上有差异),雌雄蕊原基同时出现在每一个花器官分化过程中,但是,可育性结构部分的形成取决于其原基是否分化成所应有的结构:雄蕊原基分化形成花药与花丝,雌蕊原基分化形成花柱、柱头和子房。臭椿单性花的形成是由于两性花中雌蕊原基的退化所造成,其机理有待于进一步研究。     

Flower structure and sex expression of leaky dioecy in Eurya obtusifolia

The genus Eurya is described as strictly dioecious, but rare leaky plants have been found in some species, causing much confusion about the sex expression of the genus. Through field investigations from 2009 to 2011, morphologically hermaphroditic flowers in Eurya obtusifolia were discovered. In order to understand the sex expression of these plants, their stamens, pistils and ovules were carefully observed. Staining methods were used to assess the functional gender. The flowers of the leaky plants can be classified into six types: pistillate, staminate, cryptic pistillate, cryptic staminate, hermaphrodite and infertile. Leaky plants usually exhibit a combination of different kinds of flowers, making them either gynoecious, androecious, gynomonoecious or monoecious. Some individuals have infertile flowers. Bagging experiments verified the bisexual function of E. obtusifolia plants, some of which possess the ability to self‐pollinate. The flower morphology of leaky plants varied more than that of dioecious ones, and the number of ovules were significantly negatively correlated with the number of stamens. These plants show lower fitness than normal dioecious plants. This may owe to allocation tradeoffs or sexual genome conflicts. As leaky dioecy is rarely reported in this genus, E. obtusifolia is an important species to study in order to better understand the ecological adaptations and evolutionary pathways that lead to dioecy in Eurya. Our findings provide some evidence that dioecy of Eurya evolved from hermaphroditism, but further studies are still needed.     

Observations of inflorescence patterns and flower sex in a population of Sagittaria papillosa Buch. (Alismataceae)

Sagittaria papillosa Buch. is monoecious with unisexual flowers, pistillate below, staminate above, typically with an unbranched scape. A large population with unusual numbers of staminate and bisexual flowers on the lowest whorl of the inflorescence and many particles was quantitatively evaluated. First-formed inflorescences had more staminate and bisexual flowers than those produced later. Branched scapes were predominantly found to be the second inflorescence produced by a given plant. Genetic crosses between flowers on recemes and panicles produced no branched inflorescences. When grown under greenhouse conditions all tested plants had racemes with pistillate flowers in the lower whorls and staminate ones above. Data from soil parameters, daylengths and air temperatures are compared to reported information on modification of flower sexuality by these factors.     

MORPHOLOGY,VASCULAR ANATOMY AND EMBRYOLOGY OF PISTILLATE AND STAMINATE FLOWERS OF ASPARAGUS OFFICINALIS

Flowers of three pistillate (female), two heterogametic staminate (male) and two homogametic male genotypes of Asparagus officinalis L. were compared for morphology and vascular anatomy of the flower and for embryological development to the stage of mature ovules and pollen. Flowers are liliaceous, the staminate with rudimentary pistils and the pistillate with collapsed anthers. The uncomplicated vascular pattern differs between staminate and pistillate flowers only in the size and degree of maturation of bundles to stamens and carpels. Longer styles appear to be correlated with a greater extent of ovule development in ovaries of staminate flowers. Microsporogenesis in males is normal with wall development corresponding to the Monocotyledonous type. The tapetum is glandular and binucleate, cytokinesis successive, the tetrads isobilateral or occasionally decussate, and the mature pollen grain two-celled. A pair of heteromorphic, possibly sex, chromosomes was observed in heterogametic male plants. Anther development is initially the same in pistillate flowers, but the tapetum degenerates precociously followed by collapse of microspore mother cells. In pistillate flowers the ovules are hemitropous, bitegmic, and slightly crassinucellate. Megasporogenesis-megagametogenesis conforms to the Polygonum type. In staminate flowers ovule development is like that in pistillate flowers until degeneration starts in nucellar and integumentary cells at the chalazal end. Ovules in both homogametic male genotypes rarely complete meiosis, while in the heterogametic males it is normally completed with about one ovule in 20 flowers forming a mature megagametophyte. Since manipulation of sex expression in Asparagus could be important in developing inbred male and female lines for breeding purposes, those aspects of the morphological and embryological observations presented which might be useful in planning experiments to induce sex changes are discussed briefly.     

麻疯树雌雄花中MADS-BOX基因的表达分析

麻疯树(Jatropha curcas)种子含油率高,种子中的油类物质可作为生物柴油被开发和利用,是极具潜力的生物质能源树种之一。麻疯树雌雄异花,在自然条件下雄花数量通常远远大于雌花,这大大限制了种子和油的产量,因此开展麻疯树性别分化与花发育分子机理的研究具有重要意义。该研究选取10个麻疯树的MADS-BOX基因(JcAGL1,JcAGL6,JcAGL9,JcAGL11,JcAGL15,JcAGL61-3,JcAGL62-1,JcAGL62-6,JcAGL62-7,JcAGL80-2),提取麻疯树早期发育各个阶段的雌雄花总RNA,并反转录成cDNA,采用实时荧光定量方法,探索早期发育不同阶段的麻疯树雌雄花目的基因的表达情况。结果表明:目的基因在发育起始的雌雄花中的表达具有差异,比如JcAGL6和JcAGL15在雄花中表达量要高于雌花,而JcAGL1,JcAGL9和JcAGL11在雌花中的表达量要高于雄花,这说明花原基中目的基因表达会直接或间接决定性别分化的方向;在之后的发育过程中,目的基因的表达情况在雌雄花中有所不同:随着花的发育,目的基因在雌雄花中的表达量变化存在差别,这反应出麻疯树雌雄花发育中目的基因表达模式上的差异;另外,也能看出在此过程中各个目的基因又发挥着不同的功能。该研究结果为进一步探究麻疯树雌雄花发育相关基因的表达提供了理论依据,为了解麻疯树性别分化和花发育的分子机理奠定了基础。     

Unisexual flower, spikelet, and inflorescence development in monoecious/dioecious Bouteloua dimorpha (Poaceae, Chloridoideae)

Unisexual flowers have evolved repeatedly in the angiosperms. In Poaceae, multiple transitions from bisexual to unisexual flowers are hypothesized. There appear to be at least three distinct developmental mechanisms for unisexual flower formation as found in members of three subfamilies (Ehrhartoideae, Panicoideae, Pharoideae). In this study, unisexual flower development is described for the first time in subfamily Chloridoideae, as exemplified by Bouteloua dimorpha. Scanning electron microscopy (SEM) and anatomy were used to characterize the development of male (staminate) and female (pistillate) flowers, spikelets, and inflorescences. We found the developmental pathway for staminate flowers in B. dimorpha to be distinct from that described in the other three subfamilies, showing gynoecial arrest occurs at a different stage with possible loss of some cellular contents. However, pistillate flowers of B. dimorpha had some similarity to those described in other unisexual-flowered grasses, with filament and anther differentiation in abortive stamens. Comparing our findings with previous reports, unisexual flowers seem to have evolved independently in the four examined grass subfamilies. This analysis suggests the action of different genetic mechanisms, which are consistent with previous observations that floral unisexuality is a homoplasious condition in angiosperms.     

倒地铃花的形态分化与花药结构研究

利用体视显微镜、半薄切片和超薄切片法对倒地铃(Cardiospermum halicacabum Linn.)雄花和假两性花开花过程及花药发育过程进行了观察和比较研究。结果显示:(1)花蕾发育早期,倒地铃雄花和假两性花的花蕾形态没有区别;花蕾发育后期,雄花雌蕊退化,假两性花雌蕊继续发育,花蕾外部形态出现差异;开花时雄花花药开裂,假两性花花药不开裂。(2)倒地铃雄花和假两性花均具四室花药,呈蝶形;花药壁细胞从外到内依次是表皮、药室内壁、中层(2层)和绒毡层;花药壁发育为基本型,绒毡层为单核分泌型,四分体为四面体型,花粉粒两核;开花时雄花和假两性花中层都有残留;小孢子液泡化时,绒毡层开始降解,两核花粉粒时,假两性花绒毡层降解较快。(3)雄花药室内壁次生加厚完全,裂口区发育,连接同侧花粉囊的连接组织降解,花药开裂;假两性花药室内壁次生加厚不完全,具唇形细胞,药隔细胞壁未降解,同侧花粉囊未连通,花药四室,不开裂;假两性花成熟花粉粒细胞质稀少,内壁不完整。本研究结果表明,倒地铃的雄花是由两性花在发育早期雌蕊停止发育形成的,假两性花则由两性花在发育晚期雄蕊功能退化造成的。     

Quantitative developmental analysis of homeotic changes in the inflorescence of Philodendron (Araceae)

Background and Aims: The inflorescence of Philodendron constitutes an interestingmorphological model to analyse the phenomenon of homeosis quantitativelyat the floral level. The specific goals of this study were (1)to characterize and quantify the range of homeotic transformationin Philodendron billietiae, and (2) to test the hypothesis thatthe nature of flowers surrounding atypical bisexual flowers(ABFs) channel the morphological potentialities of atypicalbisexual flowers. Methods: Inflorescences of P. billietiae at different stages of developmentwere observed using SEM. The number of appendices in male, femaleand sterile flowers were counted on 11 young inflorescences(5–6 flowers per inflorescence). The number of staminodesand carpels on ABFs were counted on 19 inflorescences (n = 143).These data were used for regression and ANOVA analyses. Results: There was an average of 4·1 stamens per male flower,9·8 carpels per female flower and 6·8 staminodesper sterile male flower. There was an average of 7·3floral appendices per atypical flower. Staminodes and carpelsare inserted on the same whorl in ABFs. A negative exponentialrelationship was found between the average number of staminodesand the number of carpels in ABFs. If only the ABFs consistingof less than six carpels are considered, there is a linear relationshipbetween the number of carpels and the average number of staminodes.The value of the slope of the regression equation indicatesthat on average, in P. billietiae, 1·36 carpels are replacedby one staminode. Conclusions: In P. billietiae, the number of appendages in female flowersimposes a constraint on the maximum total number of appendages(carpels and staminodes) that can develop on ABFs. The quantitativeanalyses indicate that the average number of different typesof floral appendages on an ABF and the number of organs involvedin a homeotic transformation are two independent phenomena.     

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。