THE EFFECTS OF HORMONES UPON THE DEVELOPMENT OF EXCISED FLORAL BUDS OF AQUILEGIA

Young excised floral buds of Aquilegia were grown on defined medium containing kinetin, indoleacetic acid (IAA), or gibberellic acid (GA₃). Only when 10⁻⁶ or 10⁻⁷ m kinetin was added to the basal medium was there a significant increase in the number of initiated whorls of primordia. Buds on the basal medium or on medium with IAA or GA₃ failed to initiate carpels. On medium with 10⁻⁶ or 10⁻⁷ m kinetin, buds successfully initiated a normal whorl of five carpels. A high level of inorganic nitrogen was also required for the initiation of carpels. With 10⁻⁵ m kinetin, individual buds initiated from 6–18 carpels. Staminodial primordia of these buds were replaced with carpels, or the floral apex enlarged to accommodate a single whorl of many carpels. Kinetin did not support the further differentiation of the floral organs. Sepals, petals, and carpels did differentiate on medium with GA₃, but stamens aborted. However, on medium with GA₃ and kinetin, stamen primordia differentiated into short filaments and anthers. Further unknown growth factors appear to be required for the complete differentiation of floral primordia into mature organs.     

Effect of plant population on ear differentiation and growth in maize

Maize was grown at 2.47, 3.70 and 4.94 plants m^-2. Ears were apparently initiated about 2 days later in the densest than in the least dense population, but rate of formation and final number of female floret primordia per row of the ear were similar in the two populations. In the densest population fewer primordia produced fully developed florets than in the least dense population and of those that were produced fewer extruded silks before pollen shedding ceased. Relative growth rates of dry mass and of length of the ear before flowering were unaffected by density, but with the apparent later start of ear growth in the densest population ears were lighter and shorter than in the least dense population. Number of florets per row of the ear at the time of flowering decreased slightly with increase in population. Number of kernels per row decreased with time after flowering, especially in the densest population. At final harvest size and mass per kernel, number of kernels per ear, length and circumference of the rachis, and space per kernel on the rachis all decreased with increase in population. Thinning the intermediate population at various times from shortly before flowering until near final harvest increased mass of grain and shoot, and number of kernels of the remaining plants at final harvest. The extent of the increases decreased progressively the later the thinning. Mass and volume per kernel were increased by thinnings done up to the time of flowering, but were unaffected by thinning from c. 16 days after flowering.     

FLOWER DEVELOPMENT IN DIOECIOUS SPINACIA OLERACEA (CHENOPODIACEAE)

Spinacia oleracea (Chenopodiaceae) is a potential model system for studies of mechanisms of sex expression and environmental influences on gender in dioecious species. Development of the male and female flowers and inflorescences of spinach were studied to determine when the two sex types can be distinguished. We found that female inflorescence apices are significantly larger than those of the male. Flower primordia are similar in size prior to perianth initiation, but the male primordia develop at a faster rate. Another distinguishing feature at this early stage is the larger bract subtending the female primordium. The two flower types become readily distinguishable when the perianth initiates. Male flowers produce four sepals and four stamens in a spiral pattern in close succession. Female flowers produce two alternate perianth parts that enlarge somewhat before the gynoecium becomes visible. There are no traces of gynoecia in male flowers or of stamens in female flowers. We propose that plant sex type is determined before inflorescence development, prior to or at evocation.     

FLOWER CULTURE OF A MALE STERILE STAMENLESS-2 MUTANT OF TOMATO (LYCOPERSICON ESCULENTUM)

Young floral buds of a male sterile stamenless-2 (sl-2/sl-2) mutant of tomato were cultured, at the sepal primordia stage, in a liquid Murashige and Skoog medium containing either benzylaminopurine (BAP) or gibberellic acid (GA₃) or both. In the basal medium (BM), the buds initiated petal and stamen primordia only and they showed limited development. In buds grown in BM supplemented with 10^–6 M BAP, all types of organ primordia were initiated but the petals remained small and the stamens and carpels were immature. Well-developed flowers with a normal complement of floral organs were, however, produced in a medium containing both BAP (10^–6 M) and GA₃ (10^–7 M to 10^–5 M). The development of stamens was variable and ranged from the complete absence of microsporogenesis to the formation of abnormal pollen. Gynoecium development was normal and ovules with megaspores were produced in the ovary. The results show that male sterility in the sl-2/sl-2 mutant can be expressed in vitro and that GA₃ is essential for the in vitro growth and development of all the floral organs of this mutant.     

ORGAN INITIATION AND THE DEVELOPMENT OF UNISEXUAL FLOWERS IN THE TASSEL AND EAR OF ZEA MAYS

The development of the unisexual male and female flowers of Zea mays from bisexual initials in both tassels and ears has been reinvestigated with SEM and TEM. The early stages of spikelet branch primordia, spikelet initiation, and early flower development are similar in both flowers, though differences in rates of growth of glumes, lemmas, and palea were detected. In both tassel and ear flowers, a pair of stamens arises opposite the lemmas and a third stamen initiates later at right angles to the first pair but from a point on the meristem below its insertion. Gynoecia develop on both tassel and ear flowers first as a ridge which overgrows the apical meristem giving rise to the stylar canal and the elongate silk. Male flowers arise in the tassel through selective vacuolation and abortion of the cells of the early gynoecium. The single female flower in each ear spikelet arises through the vacuolation and abortion of stamens in the upper flower and the repression of growth of and the eventual regression of the lower flower in each spikelet. The significance of these selective organ abortions for practical applications is discussed.     

Comparative floral development in male and female plants of Palmer amaranth (Amaranthus palmeri)

Premise

Characterizing the developmental processes in the transition from hermaphroditism to unisexuality is crucial for understanding floral evolution. Amaranthus palmeri, one of the most devastating weeds in the United States, is an emerging model system for studying a dioecious breeding system and understanding the biological traits of this invasive weed. The objectives of this study were to characterize phases of flower development in A. palmeri and compare organogenesis of flower development in female and male plants.

Methods

Flower buds from male and female plants were dissected for light microscopy. Segments of male and female inflorescences at different stages of development were cut longitudinally and visualized using scanning electron microscopy.

Results

Pistillate flowers have two to three styles, one ovary with one ovule, and five obtuse tepals. Staminate flowers have five stamens with five acute tepals. Floral development was classified into 10 stages. The distinction between the two flower types became apparent at stage four by the formation of stamen primordia in staminate flowers, which developed female and male reproductive organs initially, as contrasted to pistillate flowers, which produced carpel primordia only. In staminate flowers, the putative carpel primordia changed little in size and remained undeveloped.

Conclusions

Timing of inappropriate organ termination varies across the two sexes in A. palmeri. Our study suggests that the evolution of A. palmeri from a cosexual ancestral state to complete dioecy is still in progress since males exhibited transient hermaphroditism and females produced strictly pistillate flowers.     

Sex expression in floral buds of Acorn squash

Summary Staminate flowers of the Acorn squash (Cucurbita pepo, cv. Table Queen) are initiated monosexually, pistillate flowers are initiated bisexually. Pistillate flower primordia are physiologically bisexual from a stage below 0.3 mm until they are 0.8 mm in width.Male and female floral buds were cultivated on White's medium, modified according to Galun et al. (1963). In excised female floral buds the relative development of stamens is better than in in-situ floral buds; the opposite relationship holds for ovary development. The smaller the bud at excision, the greater the stamen growth and the lesser the ovary growth, both in absolute and relative terms. Although smaller pistillate floral buds cultured in vitro showed a greater increase in maleness than larger ones, there was no difference in the percentage of female buds between explants of the two sizes when originating from the mixed stem region (that is, the stem region carrying male and female flowers). Addition of 3-indoleacetic acid and the anti-auxin p-chlorophenoxyisobutyric acid to standard medium in equal amounts resulted in somewhat better development of excised floral buds than addition of each of these compounds alone.The percentage of female buds among explants originating from identical nodes was not significantly influenced by 3-indoleacetic acid or p-chlorophenoxyisobutyric acid.Weizmann Memorial Fellow. The studies reported in this publication were mainly undertaken at the Weizmann Institute of Science     

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。     

Maturation of stamens and ovaries on cultured ear inflorescences of maize (Zea mays L.)

Observations were made on the maturation of stamens and ovaries from cultured maize (Zea mays L.) ear inflorescences. Immature ears (5.1–10.0 mm long) of maize were cultured in kinetin medium to study microsporogenesis and pollen maturation in developing stamens. Male spikelets developed on ears cultured in kinetin medium. Meiosis-I began by 7 days of culture in the developing anthers and the mature tri-nucleate pollen grains were developed by 20 days of culture. Further, kinetin was required in the culture medium for at least initial 5 days to obtain the microspores in differentiated stamens.To observe the embryosac formation in developed ovaries, ears were cultured in control, kinetin (10.1–15.0 mm long ears) medium, and kinetin + gibberellic acid (5.1–10.0 mm long ears) medium. Formation of embryosacs was noticed in the developed ovaries which were sampled after 20 days of culture. This differential flower development using two growth regulators provides an opportunity to uncover the biochemistry and physiology of micro- and mega-gametophyte development in maize.     

IN VITRO CULTURE OF FLORAL BUDS OF AQUILEGIA

Tepfer, S. S., R. I. Greyson, W. R. Craig, and J. L. Hindman. (U. Oregon, Eugene.) In vitro culture of floral buds of Aquilegia. Amer. Jour. Bot. 50(10): 1035–1045. Illus. 1963.—Floral buds at various stages of development, from early stages before sepal initiation to late stages with young carpel primordia present, were grown in culture on various agar media. A basic medium containing White's minerals, Nitsch's trace elements, coconut milk, sucrose, and assorted water-soluble vitamins was developed for growth of the buds. The addition of indoleacetic acid, gibberellic acid, and kinetin to the basic medium extended the developmental limits of buds at nearly all stages and decidedly improved the continued development of carpels. On this medium buds produce all of their organ primordia, growing from early stages to about the size of flowers at anthesis, but will not develop unless the sepals are removed. This inhibiting effect of sepals is not understood at this time. Stamen development is consistently poor beyond the point of differentiation of anther and filament, even with the addition of hormones. The development of buds in culture is illustrated and compared with development in intact plants. With further improvement of the medium, it is hoped that these buds may be used for experiments testing theories of floral morphogenesis.     

In vitro flowering and production of viable pollen of cucumber

Flowers were produced on sterile cucumber (Cucumis sativus L.) plants grown in vitro from seed and micropropagated shoots from stem fragments. The highest numbers of flowers on plants from both sources were produced on Murashige and Skoog (MS) medium without plant growth regulators (PGR), as well as with 6 μM of kinetin (Kin). Plants cultured on MS medium supplemented with 8.9 μM benzyladenine (BA) and 1.1 μM 1-naphthaleneacetic acid (NAA) did not flower. In vitro grown plants produced fewer, smaller flowers compared with greenhouse-grown plants. Male and female flowers developed on plants grown in vitro from seed and were morphologically similar to flowers on greenhouse grown plants. Micropropagated shoots produced male flowers with altered morphology. The highest viability (72.9 ± 4.2%) and germination (69.5 ± 4.1%) of pollen were observed for plants grown from seed on MS medium supplemented with 6 μM Kin. Cytological observations of meiosis in anthers of male flowers from in vitro grown plants revealed abnormalities, such as monads, dyads, triads, polyads, microcytes and degeneration of tetrads, causing reduced viability and germination of pollen. The fewest meiotic irregularities in pollen mother cells were observed in plants grown on MS medium that was PGR-free (12.1 ± 0.9%) or with 6 μM Kin (20.9 ± 1.7%).     

High Frequency Multiple Shoot Regeneration from Decapitated Embryo Axes of Chickpea and Establishment of Plantlets in the Open Environment

Multiple shoot regeneration from the cut plumular ends of embryo axes of chickpea (Cicer arietinum L.) was evaluated on Murashige and Skoog medium having different concentrations of thidiazuron (TDZ) (0.1 to 10.0 mg dm^–3) 6-benzylaminopurine (BAP) (0.5 and 1.0 mg dm^–3), kinetin (0.5 and 1.0 mg dm^–3) or zeatin (2.0 and 4.0 mg dm^–3). TDZ (0.2 mg dm^–3) was found to be the most effective cytokinin as it produced multiple shoots in 100 % of the explants from genotypes C235, ICC5166, ICC12269, ICC4951, ICC11531, BG256 and a local cultivar. Shoots were elongated on growth regulator-free medium, and rooted on growth regulator-free medium containing 1/4 MS salts + full vitamins + 3 % sucrose. Plantlets formed were acclimatized for 12 – 15 d in MS medium with a gradual reduction in sucrose concentration and transferred into pots filled with soil and kept in the field; this resulted in more than 70 % survival. The plants developed normally and produced fertile flowers and set seeds. Low temperatures, maximum 19.0 °C, and minimum 8.2 °C, during the first 15 d of transfer favoured survival on transfer to pots.     

Sex expression in monoecious cucumbers micropropagated in vitro

The effects of plant growth regulators (PRGs) on the induction of flowering and sex expression in micropropagated cucumbers are presented. The highest number of male flowers (6.0 ± 0.7 per plant) was produced by cv. Kmicic F1 on the Murashige and Skoog (MS) medium supplemented with 4.0 μM kinetin. The highest number of female flowers (3.1 ± 0.3) was also observed in cv. Kmicic F1 on either control (PRG-free) medium or medium supplemented with 6.4 μM indole-3-acetic acid (IAA). The MS medium supplemented with 4.4 μM benzyladenine inhibited flower formation. The highest percentage of flowering plantlets (67.5 ± 7.5) was observed on the control MS medium after 16 weeks of culture. Female-to-male flower ratio was influenced by the culture media and changed during cultivation. The highest pollen viability (60–70 %) was observed in anthers of plants cultured on the control medium and the medium with IAA.     

Effects of Gibberellic Acid on Floral Development in vivo and in vitro in the Cleistogamous Species, Lamium amplexicaule L.

Using established developmental markers in the corolla and androeciumof Lamium amplexicaule L., we investigated the apparent inductionof open (chasmogamous — CH) from closed (cleistogamous— CL) flowers after application of GA₃ (0.1 mM). In vivotreatment of potentially CL flower primordia caused cell expansionbut not the increased cell division in anthers and corolla necessaryto convert a CL into a CH flower. Floral primordia that appearedto be of undetermined floral type were grown in vitro on a basalmedium supplemented with kinetin (0.1 p.p.m.) and grew to maturityas CL flowers. On media additionally supplemented with GA₃,flowers underwent anthesis, but no true CH flower was produced.Gibberellins appear to be directly responsible for anthesisin the CH flower; but additional, as yet unknown growth factorsare involved in the switch from CL to CH floral form in a developinginflorescence. floral morphogenesis, Lamium amplexicaule L, henbit, cleistogamy, gibberellin, kinetin, anthesis     

THE ONTOGENY OF THE INFLORESCENCE AND FLOWER OF LIQUIDAMBAR STYRACIFLUA L. (HAMAMELIDACEAE)

A developmental study of the inflorescence of Liquidambar styraciflua L. was conducted to clarify morphological discrepancies reported in the literature. Salient features of development are: 1) the inflorescence apex results from the conversion of a terminal, vegetative apex; 2) partial inflorescence apices arise as ellipsoid structures in axils of leaves, bracts, or transitional phyllomes; 3) development of male heads is acropetal whereas female heads differentiate basipetally; 4) the partial inflorescence apex becomes segmented into several distinct subunits indicating an axillary branch system of the third order; 5) distinct individual floral primordia are initiated on the subunits; 6) a complete absence of perianth development; 7) inception of carpel primordia in flowers of lower male heads as well as female heads, but a failure of the gynoecium to develop beyond an incipient stage in male heads; and 8) development of sterile structures around the base of the styles of only female flowers near the time of anthesis. Carpellary characteristics of the sterile structures are described, their morphological nature is discussed, and the phylogenetic position of Liquidambar is evaluated.     

栝楼不同性别花芽分化形态解剖特征观察

采用体视显微镜、石蜡切片和树脂切片技术对栝楼(Trichosanthes kirilowii Maxim.)不同性别花芽分化发育时期的外部形态和内部解剖结构进行了观察。结果显示,栝楼花为雌雄异株,仅有雌花、雄花两种性别分化,且雄花的发育速度明显快于雌花的发育速度。栝楼雌雄花芽长0.2 mm左右已完成性别分化;栝楼雄花为单性花,分化过程可分为6个时期,整个发育过程仅见雄蕊原基的分化及生长。栝楼雌花为"两性花",分化过程可分为7个时期,存在雌蕊和雄蕊共同发育阶段,后期雄蕊发育败退。本研究明确了不同性别栝楼花芽发育发生的各个阶段、形态变化特点、外部形态变化特征以及雌雄花芽的分化差异,建立了雌雄花芽内部结构分化与外部形态之间相关性,为栝楼早期幼苗鉴定及性别分化研究提供了一定的参考。     

Clonal propagation of Stevia rebaudiana Bertoni by stem-tip culture

Clonal propagation of Stevia rebaudiana has been established by culturing stem-tips with a few leaf primordia on an agar medium supplemented with a high concentration (10 mg/l) of kinetin. Anatomical examination has suggested that these multiple shoots originate from a number of adventitious buds formed on the margin of the leaf. Innumerable shoots can be obtained by repeating the cycle of multiple-shoot formation from a single stem-tip of Stevia. These shoots produce roots when transferred to a medium containing NAA (0.1 mg/l) without kinetin. The regenerated plantlets can be transplanted to soil.     

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.     

Theligonum cynocrambe: Developmental morphology of a peculiar rubiaceous herb

The annual Mediterranean herbTheligonum cynocrambe shows a peculiar combination of morphological characters, e.g., switch from decussate to spiral phyllotaxis with 90–100° divergence, combined with a change from interpetiolar to lateral stipules, anemophily, lack of calyx, flowers often dimerous to trimerous, corolla fused in both male and female flowers, male flowers extra-axillary, with 2–19 stamens per flower, female flowers axillary, with inferior uniovulate ovary, basilateral style and perianth, nut-like fruits with elaiosome. In male flowers the androecium emerges as an (uneven) elliptical rim with a central depression. This common girdling primordium is divided up into several stamen primordia. In male flowers with low stamen number the stamen primordia may occupy the corners alternating with the corolla lobes. There are no epipetalous androecial primordia that secondarily divide into stamens. Male flowers occasionally show a hemispherical base that may be interpreted as remnant of the inferior ovary. In female flowers a ring primordium grows into a tube on which the petal lobes arise. The perianth and style become displaced adaxially by uneven growth of the inferior ovary. The ovary is basically bilocular. The lower region of the ovary is provided with a septum that is overtopped and hidden by the single curved ovule.Theligonum is referred to theRubiaceae-Rubioideae, with theAnthospermeae andPaederieae as most closely related tribes.     

Scanning Electron Microscopic Studies on the Organ Development of

The pattern of development of the floral parts of Longan flower was followed using scanning electron microscope. Floral initiation begins with the formation of calyx protrusions around the floral apex. After the calyx protrusions have appeared, the petal primordia at the base of the floral apex start to appear and then followed by the androecium primordia which appear at the periphery of the floral apex. Gynoecium formation begins much later (at about 30 days after floral initiation). In the male flower, androecium develops normally forming anthers and filaments. Anthers also develop in the female flowers but they are smaller and the filaments much shorter. Gynoecium in the female flower is well developed and when mature it produces a long style, a two-prong-stigma and two ovaries. In the male flower the gynoecium is poorly developed the style is short and the stigma seldom splits. Ovaries are also poorly developed in the male flower. In addition to male and female flowers, Longan also forms a number of abnormal flowers with poorly developed androecium and gyn6ecium. Male and female flowers only become apparent at about 40 days after the initiation of flower differentiation. Prior to this it is difficult to know whether a particular developing flower is going ultimately to become a male or female flower. The formation of abnormal flowers also become obvious' at about 40 days after the initiation of flower differentation.