Comparative floral ontogeny of Maesa (Maesaceae), Aegiceras (Myrsinaceae) and Embelia (Myrsinaceae): taxonomic and phylogenetic implications

The comparative floral ontogeny of five species belonging to the primuloid clade of the Ericales are investigated, viz. Maesa japonica, M. perlarius, Aegiceras corniculatum, Embelia laeta and E. ribes. All five species basically show 2/5-spiral phyllotaxis of the sepal primordia, although with some minor modification (particularly in Embelia, where the flowers are predominantly tetramerous). The phyllotaxis of the common petal-stamen primordia is also 2/5-spiral in the Maesa and Aegiceras species investigated, but appears to be unidirectional in Embelia. All five species develop common petal-stamen primordia in which the resultant petal primordia are larger than the stamen primordia, and in which the stamens develop proximally on the adaxial flank of the common primordia. Growth of the placenta in Maesa and Aegiceras partially embeds the ovules, but in Embelia the ovules are almost fully immersed in placental tissue at maturity. A comprehensive review of all previously published studies of floral ontogeny of primuloid genera is presented, and the phylogenetic significance of the variation between genera is evaluated with reference to recently published cladograms.     

Allonia decandra: Floral remains of the tribeHamamelideae (Hamamelidaceae) from Campanian strata of southeastern USA

A single flower, detached anthers with in situ pollen grains, and isolated seeds from Campanian strata (Upper Cretaceous) of Georgia, southeastern USA, document the presence of plants assignable toHamamelidaceae in the Upper Cretaceous. The fossil flower is actinomorphic, pentacyclic and pentamerous. Irregular sepals are preserved as lobes of the floral cup, and petals are narrow, with parallel margins. The androecium has two whorls of functional stamens. Anthers are tetrasporangiate, dehisce through two valves, and have strongly elongate connective protrusions which converge over the center of the flower. The organizational and architectural features of the fossil document its affinity within subtribeLoropetalinae (Hamamelideae, Hamamelidoideae). Cladistic phylogenetic analyses using parsimony were conducted to explore the relationships between the fossil flower and extant genera of the tribeHamamelideae. The strict consensus of the four most parsimonious trees showsHamamelideae andLoropetalinae as well-supported monophyletic taxa. The fossil flower is clearly included within theLoropetalinae, and is placed as sister taxon to the southeastern Asian genusMaingaya. The occurrence of fossils assignable toLoropetalinae during the Campanian documents the existence ofHamamelidaceae with a level of floral organization and character evolution equivalent to that of extant genera, early in the evolutionary history of the family.     

石山苣苔属(苦苣苔科)花形态演化及分类学意义

石山苣苔属(苦苣苔科)约30种,主要分布于我国南部的石灰岩地区.目前该属已知物种数虽少但花形态极其多样,是该科中分类最为困难的类群之一.基于分子证据,其它8个属中花形态迥异的一些物种被并入石山苣苔属.然而,该属花形态的演化趋势缺乏系统性的研究,传统分类对属的界定与分子系统学研究结果相矛盾的原因,以及是否有形态特征支持新界定的石山苣苔属还不清楚.该研究中,总共编码了19种石山苣苔属植物和9种报春苣苔属植物的35个形态特征,其中包括26个花部形态特征,在分子系统树上追踪了它们的演化路径.结果表明:无论属内还是属间,多数花部形态特征,尤其以往属的分类界定特征,在演化过程中变化频繁且发生了高度同塑性演化,这是导致传统形态分类不自然的关键因素.此外,在观察研究的所有特征中,花丝和柱头的差异可能在石山苣苔属植物共同祖先中经历了演变,或可用于区分石山苣苔属与其姐妹报春苣苔属的大多数种类.因此,在苦苣苔科植物的分类学研究中应当慎用这些花部性状作为分类依据,而且应对形态特征进行广泛地观察研究,在密集的取样和分辨率更高、更可靠的系统树上追踪它们的演化规律.更为重要的是,需要进一步研究导致复杂形态性状演化的内在分子调控机理和外在的自然选择动力,最终更加深入地理解石山苣苔属等典型喀斯特植物的演化过程和机理.     

Notes on the floral morphology in Vivianiaceae (Geraniales)

The functional floral morphology of the three genera of Vivianiaceae (= Ledocarpaceae, Geraniales), Rhynchotheca, Viviania and Balbisia, is compared. Likely pollination mechanisms are inferred from morphology and field observations. The flowers of Viviania are nectariferous and apparently zoophilous with nectar as the (primary) pollinator reward. Balbisia has pollen flowers without nectaries, its showy corolla indicates that it is also zoophilous with pollen as sole pollinator reward; bees were observed as flower visitors. One taxon (B. gracilis) may be anemophilous. Rhynchotheca has flowers without petals, with large, pendulous anthers and lacks nectaries. It shows synchronous mass flowering in its natural populations and is evidently anemophilous. A comparison with other Geraniales shows that nectar flowers with small anthers are likely the ancestral condition in Vivianiaceae. This suggests that the pollen flowers with larger anthers of Balbisia and Rhynchotheca may represent an apomorphic condition. The documentation of pollen flowers and anemophily in Vivianiaceae expands the range of known floral and pollination syndromes in Geraniales.     

Floral Spray Transformation Can Efficiently Generate Arabidopsis

In this study, floral spray and floral dip were used to replace the vacuum step in the Agrobacterium-mediated transformation of a superoxide dismutase (SOD) gene into Arabidopsis. The transgene was constructed by using a CaMV 35S promoter to drive a rice cytosolic CuZnSOD coding sequence in Arabidopsis. The transgene construct was developed in binary vectors and mobilized into Agrobacterium. When Arabidopsis plants started to initiate flower buds, the primary inflorescence shoots were removed and then transformed by floral spray or floral dip. More than 300 transgenic plants were generated to assess the feasibility of floral spray used in the in planta transformation. The result indicates that the floral spray method of Agrobacterium can achieve rates of in planta transformation comparable to the vacuum-infiltration and floral dip methods. The floral spray method opens up the possibility of in planta transformation of plant species which are too large for dipping or vacuum infiltration.     

Floral scent chemistry and pollination ecology in phytelephantoid palms (Arecaceae)

The subfamilyPhytelephantoideae comprises three genera (Ammandra, Aphandra, andPhytelephas) and seven species of dioecious palms. The floral scents ofAmmandra dasyneura, A. decasperma, Aphandra natalia, Phytelephas aequatorialis, P. macrocarpa, andP. seemannii were analyzed by gas chromatography-mass spectrometry. We studied the pollination biology ofA. natalia, P. aequatorialis, andP. macrocarpa, and tested how the synthetically produced main constituents of the floral scents ofAphandra andPhytelephas attracted insects in two natural populations ofPhytelephas. The genera are distinct in terms of floral scents.Ammandra has sesquiterpenes,Aphandra (+)-2-methoxy-3-sec-butylpyrazine, andPhytelephas p-methyl anisol. These constituents dominated the scents quantitatively and qualitatively. The similarity between scents of male and female inflorescences was 76.5% inAmmandra, 84.2% inAphandra, and >99% inPhytelephas. Different species ofAleocharinae (Staphylinidae) pollinateAphandra natalia andPhytelephas species and reproduce in their male inflorescences.Derelomini (Curculinoidae) andMystrops (Nitidulidae) only visit and pollinatePhytelephas in which male inflorescences they reproduce. A species ofBaridinae (Curculionidae) only visits and pollinatesAphandra natalia, and reproduces in its female inflorescence. The apparent reliance on one or a few floral scent constituents as attractants and few and specific pollinators may indicate co-evolution. Sympatric species ofPhytelephantoideae have different scents. We suggest that species with similar scents have allopatric distributions due to the absence of a pollinator isolation mechanism.     

Floral evolution and phylogeny in the tribeThelymitreae (Orchidaceae: Neottioideae)

The Australian orchid tribeThelymitreae, composed ofCalochilus, Epiblema, andThelymitra, is unique in theOrchidaceae because of the presence of a mitra or staminodal complex. Evidence from floral structure suggests thatEpiblema andThelymitra are sister genera and thatCalochilus is derived from aThelymitra ancestor. A Gene Pool Vortex model and a hypothetical phylogeny illustrate that introgressive hybridization, allopatry, and long distance dispersal have played a major role in the evolution of the tribe. Pollination and hybridization in the tribe are discussed with major emphasis on floral mimicry.     

Floral development and systematic position of <Emphasis Type="Italic">Arillastrum</Emphasis>, <Emphasis Type="Italic">Allosyncarpia</Emphasis>, <Emphasis Type="Italic">Stockwellia</Emphasis> and <Emphasis Type="Italic">Eucalyptopsis</Emphasis> (Myrtaceae)

We have investigated the floral ontogeny of Arillastrum, Allosyncarpia, Stockwellia and Eucalyptopsis (of the eucalypt group, Myrtaceae) using scanning electron microscopy and light microscopy. Several critical characters for establishing relationships between these genera and to the eucalypts have been determined. The absence of compound petaline primordia in Arillastrum, Allosyncarpia, Stockwellia and Eucalyptopsis excludes these taxa from the eucalypt clade. Post-anthesis circumscissile abscission of the hypanthium above the ovary in Stockwellia, Eucalyptopsis and Allosyncarpia is evidence that these three taxa form a monophyletic group; undifferentiated perianth parts and elongated fusiform buds are characters that unite Stockwellia and Eucalyptopsis as sister taxa. No floral characters clearly associate Arillastrum with either the eucalypt clade or the clade of Stockwellia, Eucalyptopsis and Allosyncarpia.We gratefully acknowledge Clyde Dunlop and Bob Harwood (Northern Territory Herbarium) for collecting specimens of Allosyncarpia, and Bruce Gray (Atherton) for collecting specimens of Stockwellia. The Australian National Herbarium (CANB) kindly lent herbarium specimens of Eucalyptopsis for examination. This research was supported by a University of Melbourne Research Development Grant to Andrew Drinnan.     

Floral development in Adonideae (Ranunculaceae)

Floral development and floral phyllotaxis in species of Adonis, Callianthemum, and Trollius (Ranunculaceae) were studied with scanning electron microscopy. The floral organs are initiated in spiral sequence and the flowers have spiral phyllotaxis. The sepal primordia are broad, crescent-shaped, and truncate, but those of petals, stamens, and carpels are rather hemispherical. A relatively long plastochron appears to be present between the last sepal and the first petal as compared with the short and equal plastochrones of all subsequent floral organs. Maturation of the stamens within the androecium appears to be centripetal. The carpels have a short ascidiate zone. Placentation is uniformly lateral, even in Adonis and Callianthemum, which have only one fertile ovule per carpel (versus median in other genera of Ranunculoideae with a single fertile ovule). In Adonis and Callianthemum at the tip of the carpel the ventral slit is gaping and the stigma is broadly exposed, whereas in Trollius the stigma is narrower and more pronouncedly decurrent along the ventral slit. The petals in Callianthemum and Trollius are more conspicuously delayed in development than those in Adonis as compared with sepals and stamens. A short carpel stipe is formed early in Callianthemum but later in Adonis and Trollius. In Trollius farreri (commonly having only five carpels in contrast to other species of Trollius) the carpels form a single (spiral) series. Thus floral development is similar in all three genera and, at a lower level, Adonis and Callianthemum are especially close but have different autapomorphies, which reflects the current classification of the genera.     

Taxonomic studies ofAsarum sensu lato

The floral vascular anatomy of 12 species representing each ofAsarum s. str.,Asiasarum, Geotaenium, Heterotropa andHexastylis are compared to clarify intergeneric relationships. The five genera have basically similar structures in floral morphology and vasculature, and consistently have a six-carpelled compound ovary and the associated similar placental vasculature. They show, however, a significant difference in the position and the constituent of the “ventral” carpellary bundles in the placental axis betweenAsiasarum-Heterotropa-Hexastylis andAsarum-Geotaenium. InAsiasarum, Heterotropa andHexastylis the ventral bundles of each carpel are basically free and antilocular as expected in the least specialized compound ovary of angiosperms; in contrast, inAsarum andGeotaenium the ventral carpellary bundles are antiseptal and heterogenous (i.e., formed by the lateral fusion of ventral bundles of adjacent carpels). Shared probable apomorphic floral vasculature, as well as shared single style-column, suggests the closest mutual relationships betweenAsarum andGeotaenium. In terms of floral morphology and anatomy,Asiasarum, Heterotropa andHexastylis retain plesiomorphies. Possible chromosomal evolution in the related genera is also discussed.     

Comparative floral morphometrics in day-flowering, night-flowering and self-pollinated Caryophylloideae (Agrostemma, Dianthus, Saponaria, Silene, and Vaccaria)

Morphological variation of flowers with different pollination modes was studied in 53 species representing five genera (Agrostemma, Dianthus, Saponaria, Silene s.l., Vaccaria) of the subfamily Caryophylloideae. All species were classified a priori as either diurnal, nocturnal, or selfing. Canonical discriminant analysis (CDA) of 13 floral characters revealed significant correlations between floral morphometry and pollination modes. A CDA of taxonomical groups represented with more than three species (Dianthus, and subgroups of Silene s.l.: Lychnis, Silene s.str., and Viscaria), revealed that the discrimination between these four taxa, based on the same floral characters, is also well supported. Main factors for the discrimination of the species with different pollination modes were characters that (a) define nectar accessibility (calyx length, calyx tooth length), (b) are related to the positioning of anthers and stigmatic areas in relation to pollinators and (c) are important for the visual attractiveness of flowers (plate width). The functional distance between the nectar source (anthophore base) and contact zone with pollinators (style tips), given as the sum of anthophore length, ovary length, and style length (hence called AOS-complex) is better correlated with the calyx length than the single characters. Further, the total AOS-complex length differs significantly between pollination modes suggesting that these characters form a functionally linked complex that is related to the pollen placement on, and stigma contact with, the pollinator's body. However, the contribution of anthophore and style length to the total AOS-complex differed significantly between Silene s.l. and other taxa indicating that the taxonomic groups follow different evolutionary ways for the construction of the functionally linked AOS-complex.     

Apple has two orthologues of FLORICAULA/LEAFY involved in flowering

Two orthologues of FLORICAULA/LEAFY, AFL1 and AFL2 (apple FLO/LFY), were isolated from the floral buds of apple trees. Their expression was detected in various tissues and during differentiation of the floral buds. Furthermore, the flowering effectiveness of each gene was assessed with transgenic Arabidopsis. Both AFL1 and AFL2 showed high homology to each other (90%) and a high degree of similarity to PTLF and PEAFLO (70%), which are homologues of FLO/LFY from poplar and pea, respectively. RNA blot analysis showed that AFL1 was expressed only in the floral bud during the transition from vegetative to reproductive growth, whereas AFL2 was expressed in vegetative shoot apex, floral buds, floral organs and root. Genomic Southern analysis showed that apple had other homologues in addition to AFL1 and AFL2. The transgenic Arabidopsis with over-expressed AFL2 showed accelerated flowering and gave rise to several solitary flowers from rosette axils directly. AFL1 had similar effects, but the phenotypes of the transgenic Arabidopsis with AFL1 were weaker than those with AFL2. These results suggest that both genes are involved in flower differentiation in apple.     

Pollination Processes in Idiospermum australiense (Calycanthaceae), an arborescent basal angiosperm of Australia’s Tropical Rain Forests

Idiospermum australiense (Diels) S.T. Blake, a rainforest canopy tree restricted to a few small populations in northeast Australia, is the only southern hemisphere representative of the Calycanthaceae. Pollination processes in Idiospermum were investigated. Flowers are protogynous, with some populations of the species being andromonoecious, whilst others are hermaphrodite. Over their 10 – 16 day floral lifespan, movements of floral organs enforced spatial and temporal separation of male and female floral function. Changes in colour and intensity of fragrance may also influence their attractiveness to potential pollinators. Pollen is present in large quantities, and appears to act as a reward for floral visitors. Insect trapping was carried out both within flowers, and immediately adjacent to inflorescences. A wide variety of arthropods was trapped, with thrips being the most abundant. Other arthropods were trapped, including fourteen species of beetle. For arthropods other than thrips, the mean numbers of adults trapped on flowers remained more or less constant throughout the floral lifetime. However, there was a significant interaction between floral age and the number of pre-adult arthropods present – as the flower aged, the mean number of larvae and eggs per flower increased. The flowers appear to act as sheltered mating and brooding sites for small insects.     

Number of floral organs in Circaeaster agrestis (Circaeasteraceae) and possible homeosis among floral organs

98.9% of 5092 flowers from 1041 individuals of Circaeaster agrestis have five floral organs, the formula is P₃A₁G₁ (73.13%), P₂A₂G₁ (25.59%), and P₂A₁G₂ (0.22%). Only 0.4% of the flowers have six floral organs and the formula is P₃A₁G₂ (20 flowers) or P₃A₂G₁ (one flower). All these flowers have one vascular bundle in the pedicel and were considered to be normal ones. There are 33 flowers (0.65%) with six or more floral organs and two vascular bundles in the pedicel and we found traces of fusion of different degree of two flowers into one. These flowers were considered as abnormal. Therefore the normal number of floral organs of C. agrestis is five and occasionally six, and the floral formulas are P₃A₁G₁ or P₂A₂G₁, sometimes P₂A₁G₂, and occasionally P₃A₁G₂ or P₃A₂G₁. A tepal in P₃A₁G₁ may be replaced by a stamen in P₂A₂G₁ or by a carpel in P₂A₁G₂ or in reverse. A carpel in P₃A₁G₂ may be replaced by a stamen in P₃A₂G₁ or in reverse. We hypothesize that there are two possibilities for the number of the floral organs to be five (six), the result of reduction from P₃A₂G₂, or there exists homeosis among floral organs.     

Separation of AG function in floral meristem determinacy from that in reproductive organ identity by expressing antisense AG RNA

The Arabidopsis floral homeotic gene AGAMOUS (AG) is a regulator of early flower development. The ag mutant phenotypes suggest that AG has two functions in flower development: (1) specifying the identity of stamens and carpels, and (2) controlling floral meristem determinacy. To dissect these two AG functions, we have generated transgenic Arabidopsis plants carrying an antisense AG construct. We found that all of the transgenic plants produced abnormal flowers, which can be classified into three types. Type I transgenic flowers are phenocopies of the ag-1 mutant flowers, with both floral meristem indeterminacy and floral organ conversion; type II flowers are indeterminate and have partial conversion of the reproductive organs; and type III flowers have normal stamens and carpels, but still have an indeterminate floral meristem inside the fourth whorl of fused carpels. The existence of type III flowers indicates that AG function can be perturbed to affect only floral meristem determinacy, but not floral organ identity. Furthermore, the fact that floral meristem determinacy is affected in all transformants, but floral organ identity only in a subset of them, suggests that the former may required a higher level of AG activity than the latter. This hypothesis is supported by the levels of AG'mRNA detected in different transformants with different frequencies of distinct types of abnormal antisense AG transgenic flowers. Finally, since AG inhibits the expression of another floral regulatory gene AP1, we examined AP1 expression in antisense AG flowers, and found that AP1 is expressed at a relatively high level in the center of type II flowers, but very little or below detectable levels in the inner whorls of type III flowers. These results provide further insights into the interaction of AG and AP1 and how such an interaction may control both organ identity and floral meristem determinacy.     

Pollen-ovule patterns in tribeTrifolieae (Leguminosae)

Relative pollen and ovule production in the genera of the legume tribeTrifolieae is explored particularly as to howMedicago with its explosive pollination mechanism compares with its allies.Medicago produces much larger, although much fewer pollen per ovule than the other five genera in the tribe; this is interpreted as a consequence of its highly specialized, irreversible pollination mechanism, which allows only one effective exchange of pollen with pollinators.Melilotus andTrifolium produce a comparatively large quantity of pollen; this can be related to their floral characteristics requiring abundant pollen to achieve fertilization. InMedicago andTrigonella, annuals convert a higher proportion of pollen grains and ovules into seeds than perennials.     

Benzylaminopurine induces phenocopies of floral meristem and organ identity mutants in wild-typeArabidopsis plants

Arabidopsis thaliana (L.) Heynh. has been used as a model system to investigate the regulatory genes that control and coordinate the determination, differentiation and morphogenesis of the floral meristem and floral organs. We show here that benzylaminopurine (BAP), a cytokinin, influences flower development inArabidopsis and induces partial phenocopies of known floral homeotic mutants. Application of BAP to wild-type inflorescences at three developmental stages results in: (i) increase in floral organ number; (ii) formation of abnormal floral organs and (iii) induction of secondary floral buds in the axils of sepals. These abnormalities resemble the phenotypes of mutants,clv1 (increase in organ number),ap1,ap2,ap3 (abnormal floral organs) andap1 (secondary floral buds in the axils of first-whorl organs). In addition, BAP induces secondary floral buds in the axils of perianth members ofapt2-6, ap3-1 andag mutants, and accentuates the phenotype of theapt2-1 mutant to resemble theapt2-6 mutant. These observations suggest that exogenous BAP suppresses the normal functioning of the genes for floral meristem identity and thereby affects flower development and the later stages of floral organ differentiation.Abbreviations BAP N6-benzylaminopurine - CK cytokinin     

Aspects of floral structure and phenology in the genusConophytum (Mesembryanthemaceae)

The complex floral structure in the southern African genusConophytum (Mesembryanthemaceae; 77 spp.) is described in detail and assigned to three basic floral types, two of which can be divided into two subtypes. Correlations between structural features and phenological patterns are demonstrated and discussed in the family context as well as in relation to the systematic subdivision of the genus.     

Androecium and floral nectaries ofHarungana madagascariensis (Clusiaceae)

The mature flower ofHarungana madagascariensis (Choisy)Poir. has an androecium of five antipetalous fascicles, consisting of four stamens each. The stamen fascicles alternate with five indented nectary scales. A SEM-study of the floral development, as well as a study of the floral anatomy was carried out to understand whether the nectariferous scales represent staminodia or are receptacular in nature and consequently whether or not the androecium ofHarungana, and theClusiaceae in general, is originally diplostemonous. The five petals originate by the splitting of petal-stamen complexes. Next the upper part of each complex differentiates basipetally in four stamens. The stamens remain fascicled and are lifted on a long stalk at maturity. Five carpel primordia are initiated united in a low ringwall. The five nectary scales appear after carpel inception and develop an external morphology reminiscent of anthers. The floral anatomy reveals an independent origin of sepal median traces and common sepal lateral traces, free petal traces, stamen fascicle traces and alternating vascular tissue which supplies the nectaries. The petal-stamen complexes are the result of a retardation in petal inception, linked with the absorption of petal tissue into the stamen primordia. The development of the stamen fascicles is discussed; it is suggested that they are of a secondary nature and do not appear as a reduction from a multistaminate androecium. The external morphology and vascular anatomy of the scales speaks in favour of a staminodial nature. The comparison with some other species of theClusiaceae gives evidence of a diplostemonous ancestry of the androecium.     

Pollination of the southwestern Opuntias

This paper presents an overview of pollination systems inOpuntia sens. lat. in the southwestern United States. The floral syndrome of large, colorful, bowl-shaped, diurnal flowers is present in all of the prickly-pear species and most of the cholla (Cylindropuntia) species in this area. Pollination by medium-sized and large bees is established for some of the species with this floral syndrome and is predicted for the others. The same floral syndrome is widespread inOpuntia in other geographical areas.—Deviations from this floral syndrome occur in certain species of southwestern chollas, one of which has nocturnal disc-shaped flowers, and in several tropical American species groups or segregate genera with red hummingbird flowers. But such cases are relatively infrequent. Divergence between species with respect to floral syndrome and pollination system is not a common feature in the evolutionary pattern ofOpuntia. Pollination of North American Cacti, IV.—Previons parts of this series:Grant & Grant 1979a,Grant & al. 1979,Grant & Grant 1979b.