Flower structure and potential bisexuality in Freycinetia reineckei (Pandanaceae), a species of the Samoa Islands

In Freycinetia reineckei the staminate flower (on the staminate spikes) comprises 3 or 4 (sometimes 2) stamens and a pistillode with 2 (sometimes 4) carpellodes, and the pistillate flower (on the pistillate spikes) is formed of a pistil with 2 (sometimes 4) carpels and of 3 or 4 (sometimes 2) staminodes. This perfect floral homology, also observed in all the other species that were studied with both pistillate and staminate material, strongly suggests that the flower of Freycinetia is basically and potentially bisexual, and may explain the occasional sexual lability and bisexuality of that flower (occurrence of both pistillate and staminate inflorescences, and/or of bisexual inflorescences with bisexual flowers and/or unisexual flowers, on the same individuals) in some species, and also the frequent occurrence of bisexual spikes in this species. These may be partitioned into pistillate, staminate, mixed and sterile zones. In the pistillate zones the flowers have the same aspect and structure as the pistillate flowers. In the staminate zones the flowers generally comprise 3 or 4 (sometimes 2) stamens and a ‘semi-pistil’ some have both stamens and staminodes. The semi-pistils are intermediate between pistils and pistillodes in length, aspect and structure, but always have placentas and ovules. In the mixed zones the flowers are generally formed of a pistil and 3 or 4 (sometimes 2) stamens, and are therefore true hermaphrodite flowers; some have both stamens and staminodes. In the sterile zones the flowers comprise a semi-pistil and 3 or 4 (sometimes 2) staminodes. The staminodes are anatomically very similar to the stamens, especially in the staminate, mixed, and sterile zones, in which they exhibit a wide range of variation in length, aspect and structure. The perfect floral homology as generic character on one hand, and the occasional bisexuality both with and without bisexual flowers and other aspects of sex expression (e.g. occurrence of both pistillate and staminate shoots on the same individuals) in some species on the other hand, seem to indicate that Freycinetia is a basically monoecious, sex changing genus.     

DEVELOPMENT OF TASSEL SEED 2 INFLORESCENCES IN MAIZE

The normal pattern of maize floral development of staminate florets on the terminal inflorescence (tassel) and pistillate florets on the lateral inflorescences (ears) is disrupted by the recessive mutation tassel seed 2. Tassel seed 2 mutant plants develop pistillate florets instead of staminate florets in the tassel. In addition, the ears of tassel seed 2 plants display irregular rowing of kernels due to the development of the normally suppressed lower floret of each spikelet. The morphology of tassel and ear florets of the recessive maize mutant tassel seed 2 has been compared to those of wild-type maize through development. We have identified the earliest stages at which morphological signs of sex differentiation are evident. We find that sex determination occurs during the same stage on tassel and ear development. Early postsex determination morphology of florets in wild-type ears and in tassel seed 2 tassels and ears is identical.     

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.     

The development of pistillate and perfect florets in Xeranthemum squarrosum (Asteraceae)

The formation of capitulum inflorescence with two different types of floret is an interesting issue in floral biology and evolution. Here we studied the inflorescence, floral ontogeny and development of the everlasting herb, Xeranthemum squarrosum, using epi‐illumination microscopy. The small vegetative apex enlarged and produced involucral bracts with helical phyllotaxy, which subtended floret primordia in the innermost whorl. Initiation of floret primordia was followed by an acropetal sequence, except for pistillate peripheral florets. The origin of receptacular bracts was unusual, as they derived from the floral primordia rather than the receptacular surface. The order of whorl initiation in both disc and pistillate flowers included corolla, androecium and finally calyx, together with the gynoecium. The inception of sepals and stamens occurred in unidirectional order starting from the abaxial side, whereas petals incepted unidirectionally from the adaxial or abaxial side. Substantial differences were observed in flower structure and the development between pistillate and perfect florets. Pistillate florets presented a zygomorphic floral primordium, tetramerous corolla and androecium and two sepal lobes. In these florets, two sepal lobes and four stamen primordia stopped growing, and the ovary developed neither an ovule nor a typical stigma. The results suggest that peripheral pistillate florets in X. squarrosum, which has a bilabiate corolla, could be considered as an intermediate state between ancestral bilabiate florets and the derived ray florets.     

Evolution of flower shape in <Emphasis Type="Italic">Plantago lanceolata</Emphasis>

Plantago lanceolata produces small actinomorphic (radially symmetric), wind-pollinated flowers that have evolved from a zygomorphic, biotically pollinated ancestral state. To understand the developmental mechanisms that might underlie this change in flower shape, and associated change in pollination syndrome, we analyzed the role of CYC-like genes in P. lanceolata. Related zygomorphic species have two CYC-like genes that are expressed asymmetrically in the dorsal region of young floral meristems and in developing flowers, where they affect the rate of development of dorsal petals and stamens. Plantago has a single CYC-like gene (PlCYC) that is not expressed in early floral meristems and there is no apparent asymmetry in the pattern of PlCYC expression during later flower development. Thus, the evolution of actinomorphy in Plantago correlates with loss of dorsal-specific CYC-like gene function. PlCYC is expressed in the inflorescence stem, in pedicels, and relatively late in stamen development, suggesting a novel role for PlCYC in compacting the inflorescence and retarding stamen elongation in this wind pollinated species.     

The developmental floral morphology of Montrichardia arborescens (Araceae) revisited

The spadix of Montrichardia arborescens contains unisexual flowers without a perianth. The pistillate flowers are located in the basal portion of the inflorescence, and the staminate flowers are located in the apical portion. There is a narrow :zone between male flowers and female flowers consisting of atypical flowers. The portion of the atypical flowers facing the staminate zone exhibits staminate characters (stamens), and the portion facing the pistillate zone has an aborted gynoecium. The floral development of Montrichurdia is compared with that of Philodendron and a new interpretation of the morphology of atypical flowers of Montrichardia is proposed. Ontogenetic evidence supports relationships with Philodendron rather than Cercestis. 2001 The Linnean Society of London     

The role of endogenous growth regulators in the differentiation processes of walnut (Juglans regia L.)

The character of endogenous growth substances was investigated in developing buds, young fruits and mature walnut leaves. The relatively high content of auxins and gibberellin-like substances was found by means of bioassays in the youngest primordia of vegetative buds. The level of auxins drops in the further course of primordia transformation into the staminate catkins. The development of leaf-buds is characterized by the accumulation of inhibitory activity as revealed by theAvena bioassay, whereas the data obtained from the lettuce bioassay indicate a pronounced stimulation. The onset of terminal bud development is also accompanied by inhibitions and it is only with pistillate flower differentiation that the temporary rise in auxin level is observed. An inhibitory activity was found in these extracts using lettuce bioassay. There is a relatively high auxin level in young fruits, mature leaves and resting buds during the mid-summer period whereas the accumulation of clearcut inhibitions is signalled by the results of lettuce bioassay. The regulatory role of growth substances in differentiation may be better understood during the second year as many leaf-abnormities appear only with the outgrowing of the bud. Abnormal catkins differ in the number of florets and stamens and some even bear pistillate flowers. Fruit development is liable to deviations in the early stages of differentiation. Abnormal fruits enable us to elucidate many structural peculiarities.     

Floral developmental evidence for the systematic position of Batis (Bataceae)

Molecular phylogenies have associated Bataceae with Salvadoraceae and Koeberliniaceae in an expanded Brassicales. Despite a long taxonomic history, the knowledge of the flower of Batis is still fragmentary. The floral development of pistillate and staminate inflorescences of Batis maritima was investigated to understand homologies of floral structures and to discuss the phylogenetic position of Bataceae within the Brassicales. There has been considerable controversy in the past about the male flower, especially on the nature of the petals and the tubular structure enclosing the flower. Developmental evidence confirms that the male flower is built on a basic tetramerous bauplan and that the tubular structure is derived from four congenitally fused sepal lobes with the three anterior lobes highly reduced. The development of petals and stamens is unidirectional, and the androecium initiates the median stamens before the lateral stamens, suggesting the existence of two whorls. The pistillate flowers are reduced to the bare minimum with two transversal carpels enclosed by a bract. Partial inflorescences function as a swollen dispersal unit. The vestigial stipules probably represent colleters and are not homologous with true stipules. Several characters of Batis are reminiscent of the Brassicaceae, although a link with Salvadoraceae and Koeberliniaceae cannot be excluded.     

Reproductive developmental complexity in the African oil palm (Elaeis guineensis, Arecaceae)

Species of the palm family (Arecaceae) are remarkably diverse in their inflorescence and floral morphologies, which make them a particularly interesting group for studies of reproductive development and its evolution. Using light and scanning electron microscopy, we describe inflorescence and flower development in the African oil palm Elaeis guineensis from the initiation of the inflorescence meristem to flower maturity. In mature palms, the inflorescence develops over 2-3 years and is characterized by individual stages within which differentiation may be either relatively slow, as in the case of early inflorescence meristem development, or rapid, as in the case of flower organogenesis. The female inflorescence bears floral triads composed of single pistillate flowers flanked by two abortive staminate flowers, whereas the male inflorescence contains single functional staminate flowers. This suggests a possible evolutionary movement from an ancestral hermaphrodite inflorescence form containing fully functional floral triads to the situation of temporal dioecy observed at present. Wild type flowers are compared to those bearing an epigenetic homeotic abnormality, known as mantled, involving an alteration of the identity of the organs in the fertile and sterile androecium.     

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。     

INFLORESCENCE AND FLORAL DEVELOPMENT IN HOUTTUYNIA CORDATA (SAURURACEAE)

The inflorescence of Houttuynia cordata produces 45–70 sessile bracteate flowers in acropetal succession. The inflorescence apical meristem has a mantle-core configuration and produces “common” or uncommitted primordia, each of which bifurcates to form a floral apex above, a bract primordium below. This pattern of organogenesis is similar to that in another saururaceous plant, Saururus cernuus. Exceptions to this unusual development, however, occur in H. cordata at the beginning of inflorescence activity when four to eight petaloid bract primordia are initiated before the initiation of floral apices in their axils. “Common” primordia also are lacking toward the cessation of inflorescence apical activity in H. cordata when primordia become bracts which may precede the initiation of an axillary floral apex. Many of these last-formed bracts are sterile. The inflorescence terminates with maturation of the meristem as an apical residuum. No terminal flowers or terminal gynoecia were found, although subterminal gynoecia or flowers in subterminal position may overtop the actual apex and obscure it. Individual flowers have a tricarpellate syncarpous gynoecium and three stamens adnate to the carpels; petals and sepals are lacking. The order of succession of organs is: two lateral stamens, median stamen, two lateral carpels, median carpel. The three carpel primordia almost immediately are elevated as part of a gynoecial ring by zonal growth of the receptacle below the attachment of the carpels. The same growth elevates the stamen bases so that they appear adnate to the carpels. The trimerous condition in Houttuynia is the result of paired or solitary initiations rather than trimerous whorls. Symmetry is bilateral and zygomorphic rather than radial. No evidence of spiral arrangement in the flower was found.     

FLORAL DEVELOPMENT IN MYRISTICA (MYRISTICACEAE)

Myristica fragrans and M. malabarica are dioecious. Both staminate and pistillate plants produce axillary flowering structures. Each pistillate flower is solitary, borne terminally on a short, second-order shoot that bears a pair of ephemeral bracts. Each staminate inflorescence similarly produces a terminal flower and, usually, a third-order, racemose axis in the axil of each pair of bracts. Each flower on these indeterminate axes is in the axil of a bract. On the abaxial side immediately below the perianth, each flower has a bracteole, which is produced by the floral apex. Three tepal primordia are initiated on the margins of the floral apex in an acyclic pattern. Subsequent intercalary growth produces a perianth tube. Alternate with the tepals, three anther primordia arise on the margins of a broadened floral apex in an acyclic or helical pattern. Usually two more anther primordia arise adjacent to each of the first three primordia, producing a total of nine primordia. At this stage the floral apex begins to lose its meristematic appearance, but the residuum persists. Intercalary growth below the floral apex produces a columnar receptacle. The anther primordia remain adnate to the receptacle and grow longitudinally as the receptacle elongates. Each primordium develops into an anther with two pairs of septate, elongate microsporangia. In pistillate flowers, a carpel primordium encircles the floral apex eventually producing an ascidiate carpel with a cleft on the oblique apex and upper adaxial wall. The floral ontogeny supports the morphological interpretation of myristicaceous flowers as trimerous with either four-sporangiate anthers or monocarpellate pistils.     

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.     

The relationship between phenology of pistillate flower organogenesis and mode of heterodichogamy in Juglans regia L. (Juglandaceae)

 We investigated the degree of organogenesis completed at the end of the growing season in pistillate flowers of heterodichogamous Juglans regia, English or Persian walnut. Terminal buds from paired cultivars, one each protandrous and protogynous, chosen to represent early, midseason and late leafing walnuts, were examined by scanning electron microscopy. Results indicate that pistillate floral primordia in protandrous individuals had not progressed beyond involucre initiation during the season prior to bloom. In protogynous individuals, floral differentiation had progressed to the initiation of perianth primordia. These observations are compared with an earlier report on staminate flower differentiation in the same cultivars where a comparable, but opposite, relationship exists. We conclude that the degree of differentiation in both staminate and pistillate flowers that must be completed between the time growth resumes in the spring and anthesis is a developmental determinant of the mode of heterodichogamy in walnut. Received: 15 June 1996 / Revision accepted: 25 October 1996     

Sex expression as affected by N6-benzylaminopurine in staminate inflorescence of Luffa cylindrica

The effect of N⁶-benzylaminopurine (BA) on the sex expressionof staminate inflorescences of Luffa cylindrica was investigated.Direct application of BA to the staminate inflorescence inducedbisexual and pistillate flowers and finally caused the inflorescenceto develop into a shoot that was similar to the main shoot.Such modification in a staminate inflorescence from the proximalto the distal nodes is usually in the order of staminate flowers,bisexual flowers, pistillate flowers and foliage leaves (shoot).Pinching of the main stem also caused the inflorescence to developinto a shoot in the absence of lateral shoots. BA-induced femalenesswas strengthened when the number of leaves remaining on theplant was increased. On the other hand, application of BA tothe shoot apex of the main stem starting at the 2-leaf stagesuppressed differentiation of the flower bud on the main stem. (Received December 22, 1979; )     

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。     

Organogenesis of Fascicled ear mutant inflorescences in maize (Poaceae)

The ontogeny of staminate tassels and pistillate ears in the maize mutant Fascicled ear was examined using scanning electron microscopy. The normal pattern of inflorescence development is perturbed by the Fascicled ear mutation at the transition stage. The Fascicled ear mutation promotes the development of an abnormal transition stage axis that is both shorter and broader than the wild type. The inflorescence apical meristem then undergoes a bifurcation, and two inflorescence axes arise in place of a single axis. Each derived inflorescence apical meristem may undergo a similar perturbation sequence. This expression of the Fascicled ear mutation may be repeated one to several times, which leads to the development of a fascicled pistillate inflorescence and a fascicled central spike in the staminate inflorescence. The apical meristems of some tassel branches are also bifurcated. Subsequent organogenesis during paired-spikelet and floral development in Fascicled ear plants follows the pattern of normal maize. However, triplet spikelets are occasionally observed. The organogenic disruption by the Fascicled ear mutation that we describe will aid genetic and molecular analysis on the regulation of inflorescence development in maize and other members of the genus Zea.     

INFLORESCENCE AND FLOWER STRUCTURE IN NYPA FRUTICANS (PALMAE)

Details of organogenesis, anatomy, and some aspects of histogenesis are described for the inflorescence units and flowers of the mangrove palm, Nypa fruticans. The genus is of special interest in evolutionary studies because of its disjunct morphology and substantial fossil record. The inflorescence is an erect monopodial axis bearing 7–9 lateral branches and ending in a pistillate head. The lowest of the lateral branches bears up to six orders of branches, the next ones progressively fewer, and the uppermost is usually unbranched. Lateral branches of all orders end in thick spicate, staminate rachillae. The rachillae and the pistillate head consist of spirally inserted sessile flowers, each borne in the axil of a bract. Staminate and pistillate flowers are similar in structure. Both have three separate sepals and three separate petals, which are loosely closed in bud. Staminate flowers have no pistillodes; nor are there any staminodes in the pistillate flower. The androecium consists of a stalk bearing three anthers distally and is shown to represent three stamens with filaments congenitally fused and anthers connate by the ventral faces of the connectives. The pistillate flower has three separate carpels, which expand rapidly so that by anthesis they much exceed the perianth. Each carpel is cupulate in shape, with a two-crested distal opening, and receives ca. 150 vascular bundles, many of which may branch dichotomously. No dorsal or ventral bundles can be definitely distinguished, but a ventrally open ring of 10–12 bundles surrounding the locule matures first. Allometric growth clearly accounts for much of the morphological disjunction in the reproductive organs of Nypa contrasted with those of other palms. Resemblances to coryphoid, ceroxyloid, arecoid, and cocosoid palms are indicated by these studies. Different combinations of characters and several distinctive features justify a separate major taxonomic category for this genus within the Palmae.