COMPARATIVE STUDY OF THE FLORAL VASCULATURE IN SAURURACEAE

The floral vascular systems are compared among all six taxa of Saururaceae, including the two species of Gymnotheca which have not been studied previously. All are zygomorphic (dorsiventrally symmetrical), not radial as sometimes reported, in conformity with dorsiventral symmetry during organogenesis. Apocarpy in the two species of Saururus (with four carpels and six free stamens) is accompanied by a vascular system of four sympodia, each of which supplies a dorsal carpellary bundle, two ventral carpellary bundles, and one or two stamen traces. The level at which the ventral bundles diverge is the major difference in vasculature between the two species. The other four taxa are all syncarpous, and share some degree of stamen adnation and/or connation. The vascular systems also show varying degrees of fusion. The two species of Gymnotheca (with four carpels and six stamens) are very similar to each other; in both, the ventral traces of adjacent carpels fuse to form a placental bundle, which supplies the ovules and then splits into a pair of ventral strands. The flowers of Houttuynia cordata (with only three carpels and three adnate stamens) are sessile. Each flower is vascularized by three sympodia; the median adaxial sympodium is longer than the other two sympodia before it diverges to supply the adaxial organs. Three placental bundles also are formed in Houttuynia, but the three bundles differ in their origin. The median abaxial placental bundle diverges at the same level as the three sympodial bundles of the flower, while the other two lateral placental bundles diverge at a higher level from the median adaxial sympodium. Anemopsis californica, with an inferior ovary of three carpels, sunken in the inflorescence axis, and six stamens adnate to the carpels, has a vascular system very similar to that of Houttuynia cordata. The modular theory of floral evolution is criticized, on the bases of the known behavior of apical meristems and properties of vascular systems. The hypothesis is supported that saururaceous plants may represent a line of angiosperms which diverged very early.     

FLORAL ONTOGENY OF ILLICIUM FLORIDANUM,WITH EMPHASIS ON STAMEN AND CARPEL DEVELOPMENT

The ontogeny of the flower and fruit of Illicium floridanum Ellis, the Star Anise, was investigated. Each of 5 or 6 bracts in each mixed terminal bud subtends either a vegetative or floral bud. The solitary flowers occur in terminal or axillary positions. Each flower has 3–6 subtending bracteoles arranged in a clockwise helix. The flowers in our material have 24–28 tepals, 30–39 stamens, and usually 13 (rarely 19) uniovulate carpels. Tepals and stamens are initiated in a low-pitched helix; carpels later appear whorled, but arise successively at different levels on the apical flanks. The floral apex is high-convex in outline with a tunica-corpus configuration; it increases in height and width throughout initiation of the floral appendages. Tepals, stamens, and carpels are initiated by one to several periclinal divisions in the subsurface layers low on the apical flanks, augmented by cell divisions in the outer layers of the corpus. The carpel develops as a conduplicate structure with appressed, connivent margins. Procambium development of floral appendages is acropetal and continuous. Bracteoles, tepals, stamens and carpels are each supplied by 1 trace; the carpellary trace splits into a dorsal and an ascending ventral sympodium. The latter bifurcates to form 2 ventral bundles. The ovular bundle diverges from the ventral sympodium. Ovule initiation occurs in a median axillary position to the carpel, an unusual type of ovule initiation. The fruit vasculature is greatly amplified as the receptacle and follicles enlarge. After carpel initiation an apical residuum persists which is not vascularized; a plate meristem develops over its surface to produce a papillate structure.     

VARIATION IN STRUCTURE AMONG FRUITS OF MALOIDEAE (ROSACEAE)

Fruits of Rosaceae subfamily Maloideae are known as pomes, a fruit type unique to the Maloideae. Using light microscopy we examined the pomes of 173 species, broadly representing the taxonomic diversity of the subfamily. Significant variation occurs among pomes in retention, orientation, and texture of calyx lobes on the distal end of the fruit; density and distribution of sclereids in the flesh; heterogeneity among flesh parenchyma cells; carpel number and number of ovules per carpel; degree of connation among the carpels and adnation between ovary and hypanthium; and texture of the core. Phenetic clustering closely links congeneric fruits. Sorbus is a notable exception; fruits of the compound-leaved species form a cluster unrelated to fruits of the simple-leaved species. Fruits of Malus are also heteromorphic. Division of the Maloideae into two tribes, Crataegeae and Sorbeae, based largely on texture of the core is not substantiated. We hypothesize that the ancestral pome had five carpels, minimal connation of carpels, minimal adnation of ovaries to the hypanthium, two ovules per carpel, and a leathery core. Various adaptations for frugivore dispersal of the seeds partially explain trends in pome evolution.     

Evidence for a Factor which Prevents the Development of Parthenocarpic Fruits in the Strawberry

Observations of fruit development in the cultivated strawberry(FragariaxAnanassa Duchesne) are interpreted as showing that,in unpollinated fruits, development of fruit tissues ceasesat anthesis because of an inhibitory factor produced by thematuring carpels. It is suggested that one of the functionsof pollination, or fertilization, is to remove this inhibitionand make the tissues susceptible to stimulation by growth substancesproduced either by the parent plant or the fertilized carpels. Receptacles in which pollination is confined to a single carpelswell only around the point of insertion of that carpel, andno swelling occurs beneath neighbouring unfertilized carpels.Likewise, after treatment of the plant with gibberellic acidswelling of fruit tissue occurred only at the base and did notextend into areas occupied by unfertilized carpels. Parthenocarpic fruits were produced in two ways; both dependedon the sup-pression of carpel development on the receptacle.In one case, modification of the plants' environment resultedin the production of leafy bracts in place of carpels. In theother, carpel development was specifically inhibited by applicationsof maleic hydrazide at the time of carpel formation. When carpels were only partially suppressed swelling occurredonly in areas either free of carpels or occupied by fertilizedcarpels, never in areas occupied by unfertilized carpels.     

Intercarpellary growth of pollen tubes in the extragynoecial compitum and its contribution to fruit set in an apocarpous species, Schisandra sphenanthera (Schisandraceae)

? Premise of the study: Apocarpous plants possess carpels that are separated in the gynoecium. Extragynoecial compita, commonly occurring in basal angiosperms, have been proposed to have the potential to increase offspring quantity in apocarpous species through the intercarpellary growth of pollen tubes. To date, the impact of an extragynoecial compitum on fruit or seed set has not been studied in any species. This study investigated the pollen tube pathway between adjacent carpels and its contribution to fruit set in Schisandra sphenanthera. ? Methods: We investigated the fruit set ratio in the field and collected hundreds of gynoecia at their full flowering stage. Pollinated carpel ratio and pollen tube pathway observations were performed using fluorescence optics. ? Key results: Pollen grains germinated and tubes extended along the pseudostyle surface. Some of them turned and entered the ovules at the end of the stigmatic crest, whereas others subsequently grew into neighboring carpels through promontory connections located at the base of the unfused carpels. No tubes were found growing on the surface of the receptacle. More than 24 carpels could be fertilized by pollen tubes from one carpel through hand pollination. The pollinated carpel ratio was significantly lower than the fruit set ratio under natural conditions. ? Conclusions: Pollen tubes from one carpel can easily cross in the extragynoecial compitum between the adjacent carpels of S. sphenanthera, and this intercarpellary growth of pollen tubes can significantly increase the fruit set of apocarpous species, at least in S. sphenanthera.     

SECONDARY-FRUIT ONTOGENY IN NAVEL ORANGE

Ontogeny of the secondary fruit of navel orange (Citrus sinensis [L.] Osbeck) was studied using flower buds and fruit from groves in north central Florida. The secondary fruit, or navel, develops as a whorl of secondary-carpel primordia within the primary-carpel whorl when the flower bud is 1.5 to 2.0 mm in length. A complete secondary gynoecium with fused ovary but separate styles and stigmas develops within the primary gynoecium before anthesis. The stigma and style of secondary carpels are not as distinct as those of primary carpels. Three types of tissue protrusions from the secondary fruit into primary-fruit locules were abnormal placentae, free secondary carpels, and secondary-carpel outgrowths, the last being the most common. An abscission layer is present in the central axis of the primary fruit near the base of the secondary ovary. This layer is indistinguishable from adjacent tissues prior to commencement of secondary fruit abscission. Parenchyma cells in the abscission layer, however, contain large numbers of starch grains during abscission. Secondary and primary fruit have similar sigmoid growth patterns; however, onset of the cell-enlargement stage is approximately 2 wk later in secondary fruit.     

COMPARATIVE MORPHOLOGY OF THE CARPEL IN THE ROSACEAE VI. POMOIDEAE: ERIOBOTRYA,HETEROMELES, PHOTINIA,POURTHIAEA, RAPHIOLEPIS,STRANVAESIA

The pomoid genera, Eriobotrya, Photinia, Pourthiaea, Raphiolepis, Stranvaesia, and Heteromeles, have compound inflorescences and biovulate carpels which become papery at maturity. The carpels of all of these except Heteromeles are fused with one another. There are open sutures in the carpels of Heteromeles, Photinia, Pourthiaea, and Raphiolepis, and in these four genera the extent of fusion of the ovular bundle with the wing bundle is related directly to the state of tegumentary fusion and to the extent of fusion of the carpel with the floral cup. In those species of Eriobotrya and Stranvaesia with closed sutures the integuments tend to be fused, as do the ovular and wing bundles, and the carpels are adnate with the floral cup for a considerable distance; in species with open sutures the integuments tend to be free, the ovular and wing bundles tend to be separate, and the extent of fusion of carpel with floral cup tends to be shorter. In genera with connate carpels the wing bundles of adjoining carpels may also be fused. The greatest extent of fusion occurs in Eriobotrya and Raphiolepis, in which there may also be attenuation and disappearance of the wing bundles above the region of ovular insertion and even reduction and disappearance of the carpellary margin.     

龙舌草果实和种子及其种苗发育形态学研究

采用形态观察和GMA切片相结合的方法,对龙舌草果实、种子及种苗发育的形态结构进行解剖学观察研究,探讨龙舌草个体发育中表现出的与水生环境相适应的结构特征.结果显示:(1)龙舌草由佛焰苞包被的果实通常由6心皮构成,心皮边缘不完全愈合,层片状胎座,水面形成的幼嫩果实被果柄拉入水中发育成熟;种子具毛,萌发时苗端先于根端发育,在胚根分化之前先出现单细胞的下胚轴毛,随后胚根发育为初生根,并在真叶的节处发育出不定根;果皮和叶肉内的维管束仅由少数细胞构成,且细胞分化不明显,有发达的通气道;叶表皮无角质层,细胞外凸,叶肉细胞数量少;表明龙舌草的结构特征与水生环境相适应.(2)研究还发现,龙舌草的种子无胚乳,下胚轴贮存种苗发育需要的大量淀粉粒,以及初生根和不定根均有根毛的结构,且与泽泻科的慈菇和泽泻的结构一致,证明水鳖科与泽泻科具有较近的亲缘关系.     

Female flower and fruit anatomy of Tetroncium magellanicum: implications for gynoecium evolution in the early divergent monocot order Alismatales

Female flower and fruit anatomy, including vasculature, are studied for the first time in Tetroncium (Juncaginaceae: Alismatales). Other members of Juncaginaceae (and the relatively close Maundiaceae) possess a peculiar type of gynoecium with pronounced carpel fusion via the floral centre. Their carpels are supplied by individual vascular traces and can be interpreted either as synascidiate (if viewed as horizontally inserted) or free and plicate (if viewed as obliquely inserted on an elongated receptacle). In Tetroncium, the gynoecium is tetracarpellate and clearly has a well‐developed synascidiate zone with septa formed by united flanks of adjacent carpels. The gynoecium of Tetroncium is supplied by a common ring of vascular tissue that splits into dorsal and heterocarpellary ventral (synventral) bundles, a condition that can be expected in a typical syncarpous gynoecium. The fruit is indehiscent and contains one or two seeds. The syncarpy of Tetroncium is of functional significance for fruit formation, as it allows the thin septa to be distorted, thus providing more space for the developing seed(s). The occurrence of typical syncarpy in Tetroncium provides further evidence for the highly homoplastic evolution of gynoecium characters in the early‐divergent monocot order Alismatales. Either the similarity between gynoecia of Maundiaceae and Triglochin (Juncaginaceae) is due to parallel evolution or the syncarpy of Tetroncium should be viewed as secondarily derived. In the latter scenario, fusion via the floral centre is probably a synapomorphy of core Alismatales (Helobiae) and more typical syncarpy evolved independently in several lineages, such as Scheuchzeria, Tetroncium and Butomus/Hydrocharitaceae. In total, Tetroncium differs from other Juncaginaceae in 13 structural characters, including ensiform leaves that are similar to those of Tofieldiaceae. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 712–724.     

COMPARATIVE MORPHOLOGY OF THE CARPEL IN THE ROSACEAE. III. POMOIDEAE: CRATAEGUS,HESPEROMELES, MESPILUS,OSTEOMELES

The carpels in Crataegus, Hesperomeles, Mespilus, and Osteomeles appear to constitute a morphologically related group: all have bony pits, ovules that tend to be acollateral (usually superposed), and clearly separate ovular and wing bundles, i.e., no “ventral” bundles, at the level of ovular insertion. In species whose carpels have no sutural opening, the integuments are more extensively fused with one another, the degree of intercarpellary fusion tends to be greater, and the carpels are fused with the floral cup to relatively higher levels than in those species whose carpels have a sutural opening. In the few cases in which wing and ovular bundles are adnate at the locular base (Crataegus monogyna, Mespilus, Osteomeles anthyllidifolia, O. Schwerinae), the extent of inter- and extracarpellary fusion and sutural closure is among the most advanced.     

A TEST OF SEVERAL HYPOTHESES FOR THE DETERMINATION OF SEED NUMBER IN AMELANCHIER ARBOREA,USING SIMULATED PROBABILITY DISTRIBUTIONS TO EVALUATE DATA

The number of seeds per fruit is variable within Amelanchier arborea trees. Because A. arborea flowers are five-carpellate and each carpel contains two ovules, we were able to use the pattern of seed maturation within fruits to test whether the failure of some ovules to develop into seeds is determined by mechanisms operating at the level of carpels, such as stigma-clogging, or by mechanisms operating at the level of ovules, such as ovule infertility. The presence of one-, two-, and zero-seeded carpels demonstrated that the number of ovules developing into seeds was not due entirely to carpel-level phenomena. In order to test the hypothesis of carpel independent seed development, without the assumption that all ovules have the same probability of developing into seeds, it was necessary to use simulation, since no conventional statistical models were appropriate. Analysis of this simulation allowed us to reject carpel independent processes as the only determinant of seed number. A mixed model of seed development, in which some carpels fail entirely and ovules in the remaining carpels develop equiprobably, was shown to be consistent with the data.     

Repression of the pea lipoxygenase gene loxg is associated with carpel development

A cDNA clone (loxg) corresponding to a gene repressed during carpel development has been isolated from a cDNA library of unpollinated carpels induced to grow by treatment with gibberellic acid (GA₃). The sequences of loxg cDNA and the deduced polypeptide have a high similarity with legume type 2 lipoxygenases, especially with Phaseolus lox1 (78.5% similarity at the protein level) and pea and soybean lox3 (83.6% and 85.4%, respectively). loxg expression is constant in unstimulated carpels but it decreases in carpels induced to keep growing by fertilization or hormone treatment. A similar pattern of repression was observed in lipoxygenase activity of pea and tomato carpels. In situ hybridization studies showed that loxg mRNAs are present in the endocarp and the mesocarp of pea pods; no loxg expression was detectable either in the pod exocarp or in the ovules. Loxg is also expressed in other young growing tissues, especially in flower organs. Nevertheless, the natural pattern of flower and fruit development is associated with loxg repression.     

Phylogeny of Impatiens (Balsaminaceae): integrating molecular and morphological evidence into a new classification

Impatiens L. is one of the largest angiosperm genera, containing over 1000 species, and is notorious for its taxonomic difficulty. Here, we present, to our knowledge, the most comprehensive phylogenetic analysis of the genus to date based on a total evidence approach. Forty‐six morphological characters, mainly obtained from our own investigations, are combined with sequence data from three genetic regions, including nuclear ribosomal ITS and plastid atpB‐rbcL and trnL‐F. We include 150 Impatiens species representing all clades recovered by previous phylogenetic analyses as well as three outgroups. Maximum‐parsimony and Bayesian inference methods were used to infer phylogenetic relationships. Our analyses concur with previous studies, but in most cases provide stronger support. Impatiens splits into two major clades. For the first time, we report that species with three‐colpate pollen and four carpels form a monophyletic group (clade I). Within clade II, seven well‐supported subclades are recognized. Within this phylogenetic framework, character evolution is reconstructed, and diagnostic morphological characters for different clades and subclades are identified and discussed. Based on both morphological and molecular evidence, a new classification outline is presented, in which Impatiens is divided into two subgenera, subgen. Clavicarpa and subgen. Impatiens; the latter is further subdivided into seven sections.     

Comparative morphology of the carpel in the Liliaceae: Wurmbaeae

The two genera of Buxbaum's tribe Wurmbaeae, Anguillaria and Wurmbea , have multiovulate carpels. There are deep septal indentations between the carpels of Anguillaria , but the wings of adjoining carpels are fused to solid septa in most species of Wurmbea. In Anguillaria the carpels have open sutures or prominent commissural markings; in Wurmbea the carpels generally lack these characteristics, and some species have a vascularized, columella-like axis in the centre of the pistil. In both genera there are a dorsal bundle, lateral bundles, and two placental bundles in each carpel. At the inner edge of the septum there are one or two septal bundles in Anguillaria and one or none in Wurmbea. The ovules are monotegmic, the integument and funiculus being partly fused in Anguillaria and mostly fused in Wurmbea. An obturator is present in Anguillaria but absent from most species of Wurmbea.     

Flower and fruit development in the Macaronesian endemicCeballosia fruticosa (syn.Messerschmidia fruticosa,Boraginaceae, Heliotropioideae)

The ontogeny of the flower and the fruit of the Macaronesian endemicCeballosia were investigated morphologically and anatomically by SEM and LM. The fruit does not break into four mericarps, but splits into two two-seeded carpids. Exo- and mesocarp wither after fruit-ripening and the endocarp constitutes the remaining outer wall. Within the stony endocarp tubular parenchymatic isles develop which are linked with the mesocarp. Subsequent disintegration pretends additional locules in the mature fruit. Similar pericarp formations are also found in someHeliotropium species but result from a different ontogeny. Therefore, although a close relationship ofCeballosia toHeliotropium is obvious, the taxon should be treated as a separate genus.Carpological investigations in theHeliotropioideae (Boraginaceae) 2. For part 1 seeHilger (1987).     

INFLORESCENCE AND FLOWER DEVELOPMENT IN THE PIPERACEAE III. FLORAL ONTOGENY OF PIPER

Floral development in Piper was compared between four-staminate species (P. aduncum and P. marginatum) and six-staminate species (P. amalago). All Piper species have a syncarpous gynoecium composed of three or four carpels. The floral apex is initiated by a periclinal division in the subsurface layer in the axil of a bract 40-55 μm high; initiation of the bracts occurs separately and considerably earlier. The floral primordium widens and the first pair of stamens are initiated at either side. The median anterior stamen forms next, and the median posterior later. This sequence is common to all species studied. In the six-staminate P. amalago, the last two stamens form simultaneously in lateral-anterior positions. The stamens hence arise as pairs, and symmetry is bilateral or dorsiventral. The three or four carpels arise simultaneously; they are soon elevated on a gynoecial ring by growth of the receptacle below the level of attachment of the carpels to produce a syncarpous gynoecium. The floral apex lastly produces the solitary basal ovule and is used up in its formation.     

Heat and drought determine flower female allocation in a hermaphroditic Mediterranean plant family

• In animal‐pollinated hermaphroditic species, larger and xenogamous flowers increase male‐biased resource allocation, whereas smaller and selfing flowers invest disproportionally more resources to female function. In Cistaceae, an entomophilous and hermaphroditic Mediterranean family, this pattern generally follows a phylogenetic signal. However, resource allocation to carpels is independent of phylogeny, which suggests trait divergences among closely related species during the diversification into different environmental conditions.
• We tested this hypothesis across 37 species of Cistaceae along a temperature and precipitation gradient, including semiarid, dry, subhumid and humid sites. We quantified the proportions of dry mass and nutrient investment to carpels and tested the influence of the climatic gradient and site‐specific precipitation on the interspecific variation in carpel resource allocation.
• Lowest and highest percentages of resource allocation to carpels ranged from 1.5–4.2% to 24.2–36.6%, respectively. The proportion of resources comprised in carpels significantly decreased with increasing precipitation/decreasing temperature. Thus, carpels comprised proportionally more resources under drier and hotter conditions, especially in semiarid sites.
• Our results demonstrate how the extent of climatic constraints is more important than phylogenetic relationships in determining stress‐induced differences in carpel resource allocation across species of Cistaceae in a Mediterranean environment. We suggest that allocation of proportionally more resources to carpels in drier and hotter sites lies within a strategy to deal with the most stressful conditions by means of a high reproductive effort.

     

The second intron of AGAMOUS drives carpel- and stamen-specific expression sufficient to induce complete sterility in Arabidopsis

Gene containment technologies that prevent transgene dispersal through pollen, fruit and seed are in immediate demand to address concerns of gene flow from transgenic crops into wild species or close relatives. In this study, we isolated the enhancer element of Arabidopsis AGAMOUS that drives gene expression specifically in stamens and carpels. By fusing this AG enhancer to a minimal 35S promoter fragment, two tissue-specific promoters, fAGIP and rAGIP in forward and reverse orientations, respectively, were created and fused to the GUS reporter. Transgenic Arabidopsis plants harboring either fAGIP::GUS or rAGIP::GUS displayed similar GUS expression specifically in carpel and stamen tissues and their primordial cells. To test their utility for engineering sterility, the promoters were fused to the Diphtheria toxin A (DT-A) gene coding for a ribosome inactivating protein as well as the Barnase gene coding for an extracellular ribonuclease, and tested for tissue-specific ablation. Over 89% of AGIP::DT-A and 68% of AGIP::Barnase transgenic plants displayed specific and precise ablation of stamens and carpels and are completely sterile. These transgenic plants showed normal vegetative development with prolonged vegetative growth. To evaluate the stability of the sterile phenotype, 16 AGIP::DT-A lines underwent two consecutive cutback generations and showed no reversion of the floral phenotype. This study demonstrates a simple, precise and efficient approach to achieve absolute sterility through irreversible ablation of both male and female floral organs. This approach should have a practical application for transgene containment in ornamental, landscaping, and woody species, whose seeds and fruits are of no economic value.     

Structural features of <Emphasis Type="Italic">Rhododendron luteum</Emphasis> flower

The flower of Rhododendron luteum (L.) Sweet has a pentamerous structure with radial symmetry. The anthers filament surface is covered by dense non-glandular hairs to the half of the height. The tubular anther dehisces along creating two openings in the anther-sac walls and the viscous pollen is released through two splits along the anther lobes. The pistil is pentamerous and the axial channel is filled with a mucilaginous secretion product which is continuous with the exudate on the stigma surface. The stigmatic papillae are densely packed and their exudate is stained intensively red for carbohydrates, while pollen grains are stained positively for lipids. The five-locular ovary has isomerous carpels (syncarpous gynoecium) and the ovary surface is covered by numerous, densely-packed glandular and non-glandular hairs protecting the nectar against transpiration. Numerous ovules per locule occur with one integument and a thin-walled megasporiangium. In carpels, oil cells occur sporadically as solitary idioblasts, located around the vascular bundles. Transmitting tissue cells contain a large central, electron translucent vacuole, filling most of the cell containing dark osmiophilic bodies homogenous or granular in appearance.     

CARPEL NUMBER AS A TAXONOMIC CRITERION IN PANDANUS

One of the criteria sometimes used in distinguishing species in the genus Pandanus has been the number of carpels which make up each phalange of the fruit. An investigation of the dependability of this criterion yielded results which do not invalidate its use, but its applicability is restricted. In addition a more precise method of obtaining the information required for the use of the criterion is described, involving counts of carpels-per-phalange and phalanges-per-cephalium based on whole-cephalium samples. Graphs illustrate some of the variation with respect to carpel number in certain species of the genus. A specific example of the application of the criterion is provided, as are several other examples without detailed data. Cautious use of and limited dependence on the criterion are suggested, and some cases of variation in carpel number within one species are cited. Like all other taxonomic criteria, it should be used in conjunction with all other data and its applicability evaluated for each particular taxonomic case rather than depended upon for quick solutions to taxonomic problems.