Gynoecium Structure Research Articles

Gynoecium structure and pollen tube pathway in the cactus family with emphasis on tribe Trichocereeae (Cactaceae: Cactoideae)

Abstract The structural details of the gynoecium are key to understanding the reproductive systems and successful diversification of flowering plants. However, the gynoecium morpho-anatomy in South American species of Cactaceae that evolved in the Andean region remains largely unknown. Here we selected 18 species, most of them of evolutionarily related genera of Cactaceae tribe Trichocereeae, to conduct a detailed comparative study. Observations were made using light, fluorescence, differential interference contrast and scanning electron microscopy. Most of the characters of the ovary and ovule were typical of the family, except for the nucellar beak in Echinopsis aurea and E. haematantha, here reported for the first time in cacti. We found evidence suggesting that the stigmatic surface covered with multiseriate trichomes, the semi-closed style type and the pollen tube transmitting tract are conserved characters among species of Trichocereeae; this finding may be explained by a phylogenetic conservatism of the investigated genera of the tribe. We integrated the available information about structural and histological characters of the gynoecium in the family, taking into account the current phylogenetic context of the examined genera. Our results reinforce the significance of floral anatomical traits for the systematics of Cactaceae.

Еволюційна й екологічна інтерпретація структури плодів однодольних рослин флори України

Background. Monocot plants in the flora of Ukraine embrace about 1000 species. The members of this group have various fruit types and ways of dissemination. In this review, the results of the evolutionary-morphological analysis of fruits in monocot plants of the flora of Ukraine and their structural adaptations to dispersal are presented. Gynoecium structure and morphogenetic fruit types in monocots. The features of the inner gynoecium structure and the relation between gynoecium and fruit structure were analyzed, and the main problems of fruit classification were detected. It was revealed that apocarpous fruits are often polymerous or oligomerous, composed of 3–6 one-seeded fruitlets. The species with inferior ovary often develop capsular fruits. One-seeded fruits represent the most reduced variant of the gynoecium structure and fruit wall anatomy, providing few structural traits for analysis. Ecological and functional features of fruits in monocots. The characteristics of dehiscent and indehiscent fruits, as well as adaptations to various ways of dispersal: myrmeco-, endozoo-, epizoo-, anemo-, hydro- and autochory are presented. Dehiscent fruits (follicle and capsule) reveal ventral, dorsal or double, dorsoventral dehiscence. One-seeded fruits and fruitlets are often adapted to hydrochory or anemochory. Trends of evolutionary changes of fruits in monocots. The most diverse fruits are found in low-species taxa of early monocots. Within the monocots, we can observe certain evolutionary trends: incomplete carpel fusion, the occurrence of superior one-seeded fruits and inferior many-seeded fruits. Each of these trends may be evoked by adaptations for pollination in gynoecium at the flowering stage. Conclusions. The present evolutionary morphological and ecological studies of fruits in monocot plants of the flora of Ukraine are impeded by controversial structural types of the gynoecium in some species and a scarcity of data on ways of dispersal of fruits and seeds. The precise carpological and carpoecological investigations of the monocots are desirable for many plants of the flora of Ukraine.

Flower and Spikelet Construction in Rapateaceae (Poales).

The family Rapateaceae represents an early-divergent lineage of Poales with biotically pollinated showy flowers. We investigate developmental morphology and anatomy in all three subfamilies and five tribes of Rapateaceae to distinguish between contrasting hypotheses on spikelet morphology and to address questions on the presence of nectaries and gynoecium structure. We support an interpretation of the partial inflorescence (commonly termed spikelet), as a uniaxial system composed of a terminal flower and numerous empty phyllomes. A terminal flower in an inflorescence unit is an autapomorphic feature of Rapateaceae. The gynoecium consists of synascidiate, symplicate, and usually asymplicate zones, with gynoecium formation encompassing congenital and often also postgenital fusions between carpels. Species of Rapateaceae differ in the relative lengths of the gynoecial zones, the presence or absence of postgenital fusion between the carpels and placentation in the ascidiate or plicate carpel zones. In contrast with previous reports, septal nectaries are lacking in all species. The bird-pollinated tribe Schoenocephalieae is characterized by congenital syncarpy; it displays an unusual type of gynoecial (non-septal) nectary represented by a secretory epidermis at the gynoecium base.

Gynoecium structure and development in core Caryophyllales: a matter of proportions

AbstractThis review based on a morphological and developmental perspective reveals a striking diversity in shapes and evolutionary trends in the gynoecium of core Caryophyllales that have affected the number of carpels, the formation of septa and the number of ovules. Two major developmental shifts are responsible for the diversity in gynoecial forms and are linked to the proportional development of carpellary tissue (ovary wall) and the floral apex. (1) Meristic change is caused by an expansion or reduction of the diameter of the floral apex. An expansion leads to polygyny linked with the development of more numerous small carpels; a reduction of space leads to lower carpel numbers, eventually resulting in a single carpel. (2) Different ovary shapes can be recognized at a mid-developmental stage predicting the further development of ovaries. With an equal growth of the ovary wall and floral apex, young ovaries take the shape of a salt-shaker; with more extensive development of the floral apex and delay of the ovary wall, a club-shaped ovary is formed; with faster growth of the ovary wall linked with intercalary expansion and a delayed growth of the floral apex, a (half-) inferior cup-shaped ovary develops. The different growth forms are the results of heterochronic shifts and affect the development of septa and ovule numbers. A common trend in the order implies a weakening and break-up of septa during development, leading to residual apical and basal septa and the shift to free-central and basal placentation. The club-shaped ovary is linked with an almost complete loss of septa and a reduction of the ovule number to one. The salt-shaker shape leads to ovaries with a massive placental column and several ovules. The cup-shaped ovary leads to a shift of ovules away from the floral apex. Developmental flexibility is responsible for a disconnection of carpel wall growth from ovular tissue. Subtle shifts in proportional growth lead to a high diversification of ovaries in core Caryophyllales and the establishment of predictable developmental trends. These trends clearly represent apomorphic tendencies, affecting different families of core Caryophyllales in different degrees. The ancestral gynoecium was probably pentamerous and isomerous with the other floral whorls, with ovules clearly separated from the carpellary wall and inserted on axile placentas corresponding to the central axis of the flower.

Anther and gynoecium structure and development of male and female gametophytes of Koelreuteria elegans subsp. formosana (Sapindaceae): Phylogenetic implications

Anther and gynoecium structure and embryological information in Koelreuteria and Sapindaceae as a whole remain understudied, as well as the evolution of imperfect flowers in the latter. The aims of this study were to analys in K. elegans subsp. formosana the anther and gynoecium structure and the development of male and female gametophytes in the two floral morphs of putatively imperfect flowers. Standard techniques were applied for LM and SEM. Compared to the normal anther development in staminate flowers, a delayed programmed cell death of tapetum, septum and middle layers on the onset of microspore stage result in indehiscent anthers in the functionally pistillate flowers. Orbicules are reported for the first time in Sapindaceae. Gynoecium development in functionally pistillate flowers is normal, whereas in functionally staminate ones a pistillode with degenerated ovules at the tetrad stage is formed. The pollen tube transmitting tract consists of one layer of epithelial cells with a small lumen in the style and ovary. The anatomy of the latter revealed an axile placentation and complete septum. Reproductive characters of Koelreuteria shared with other Sapindaceae taxa are a secretory binucleate tapetum, simultaneous cytokinesis, unicellular stigmatic papillae, Polygonum type megagametophyte with ephemeral antipodals, and ovules campylotropous, with funicular obturator and amphistomic micropyle. Imperfect flowers in the family are triggered by defective sporophytic tissues impacting on the normal development of microspores and megaspores, disrupting embryological events in staminate and pistillate organs at different developmental stages. Reconstructing ancestral character state under parsimony revealed that two ovules per locule, absence of obturator, presence of hypostase, and binucleate tapetum may be regarded as plesiomorphic for Sapindaceae. These analyses have provided information on the evolution of embryological characters in the family and its phylogenetic significance.

The unusual gynoecium structure and extragynoecial pollen-tube pathway in Phytolacca (Phytolaccaceae)

The fusion of carpels into a unified compound gynoecium is considered a dominant feature of angiosperm evolution and it also occurs by postgenital fusion during the gynoecium development in some apocarpous species. However, we found the reverse process, the separation of carpels from combined carpel primordia, during the development of the gynoecium in Phytolacca. Semithin sectioning and scanning electron microscopy were utilised to observe the structure and development of the gynoecia in Phytolacca acinosa and Phytolacca americana, fluorescence microscopy was utilised to observe the pollen tube growth in the gynoecia of the two species, and the topology method was applied to analyze the relationship between the gynoecium structure and pollen tube pathway. Although the gynoecia of P. acinosa and P. americana are both syncarpous, the degree of carpel fusion in the mature gynoecia of the two syncarpous species is different as a result of variant developmental processes. However, change in the degree of carpel fusion during the development of gynoecia in Phytolacca does not affect pollen tube growth because of the existence of the extragynoecial pollen-tube pathway. Thus, the change in the degree of carpel fusion in Phytolacca is primarily the result of diversification of developmental processes related to selection pressure.

Gynoecium structure and extra-gynoecial pollen-tube growth in an apocarpous species, Sagittaria trifolia (Alismataceae)

Apocarpy is regarded as an original feature obtained during the evolution of angiosperms. Compared with syncarpous plants, apocarpous plants have some adaptive disadvantages in apocarpous plants, for example, the number of offspring is lower under conditions of uneven pollen-tube distribution. However, in some apocarpous species, extra-gynoecial pollen-tube growth (EGPG) may remedy this disadvantage. We conducted micro-observations and field studies of Sagittaria trifolia, to investigate the gynoecium structure and the pathway of pollen-tube growth in the entire gynoecium. In a single-carpel pollination experiment, we found that the extra-gynoecial pollen tubes from a carpel of S. trifolia were able to fertilize approximately 13 carpels. Simulated EGPG in the entire gynoecium of S. trifolia revealed that its effect on the seed set could be divided into two stages: stage of low/high-level stigmas pollination, in which the cutoff point was about 0.1. The seed set would be markedly improved during the low-level stigmas pollination stage by EGPG when the maximum distance of extra-gynoecial pollen tubes could span three carpels, as in the present experiment. Our simulation also showed that the high pollen load could enhance the effect of EGPG on the seed set, and if the number of germinating pollen is triple the carpel number in the gynoecium, a 100% seed set rate would be obtained when approximately 50% of the stigmas are pollinated.

Auxin and cytokinin act during gynoecial patterning and the development of ovules from the meristematic medial domain.

The gynoecium is the female reproductive structure of flowering plants, and is the site of ovule and seed development. The gynoecium is critical for reproductive competence and for agricultural productivity in many crop plants. In this review we focus on molecular aspects of the development of the Arabidopsis thaliana gynoecium. We briefly introduce gynoecium structure and development and then focus on important research advances published within the last year. We highlight what has been learned recently with respect to: (1) the role of auxin in the differential development of the medial and lateral domains of the Arabidopsis gynoecium; (2) the interaction between cytokinin and auxin during gynoecial development; (3) the role of auxin in the termination of the floral meristem and in the transition of floral meristem to gynoecium; and (4) recent studies that suggest a degree of evolutionary conservation of auxin mechanisms during gynoecial development in other eudicots.

The gynoecium structure in Dracaena fragrans (L.) Ker Gawl., Sansevieria parva N.E. Brown and S. trifasciata Prain (Asparagaceae) with special emphasis on the structure of the septal nectary

In the gynoecium of <em>Dracaena fragrans</em>, <em>Sansevieria</em> <em>parva </em>and <em>S. trifasciata</em>, the vertical zonality of the ovary, the structural zonality of the gynoecium following Leinfellner, and the zonality of the septal nectary were studied. The ovary structure is characterised by a high parenchymatous ovary base and ovary roof as well as a long septal nectary that can be extended in both of them and opens with secretory nectary splits. The gynoecium of these species has a short synascidiate zone, a fertile hemisynascidiate zone with a median ovule attached, a hemisymlicate zone (only in <em>D. fragrans</em>) and an asymplicate zone (with postgenitally fused carpels) that comprises the ovary roof, common style and stigma. In the septal nectary, we detected three vertical zones: the basal zone of the distinct nectary in the ovary base or/and the synascidiate zone, the zone of the common nectary (in the hemisynascidiate and hemisymlicate zones) and the zone of the external nectary (the nectary splits in the asymplicate zone). The gynoecium structure in the studied species shows differences in the length of the gynoecium and septal nectary zones and also in the interrelationships of all these three types of vertical zonality.

CLV1 interacts with UFO in modulation of gynoecium development in Arabidopsis thaliana

The gynoecium is the female reproductive structure in flowering plants. It has been known that the CLAVATA1 (CLV1) gene regulates the floral meristem maintenance and floral organ number in Arabidopsis thaliana. However, whether CLV1 plays roles in gynoecium development remains uncharacterized. In this study, we showed that CLV1 genetically interacts with Unusual Floral Organs (UFO) to regulate the development of Arabidopsis gynoecium. Through forward genetic screen, we identified a clv1 allele, clv1-9, showing abnormal gynoecium development. Some epidermal cells on the growing inner surfaces of the clv1-9 gynoecium failed to fuse after contacting with each other, suggesting that CLV1 is essential for proper postgenital fusion. Furthermore, the ufo clv1-9 double mutant exhibited an expanded upper gynoecium, narrowed lower part of the ovary, altered vascular patterning and abaxial outward proliferation of the transmitting tract tissues. The transcription level of the ETT gene, which is required for gynoecium structure and pattern formation, was down-regulated in the ufo clv1-9 double mutant genoecia. Taken together, our results indicate that CLV1 functions together with UFO to regulate gynoecium development likely through modulating the expression of ETT.

Морфологія та васкулярна анатомія квітки Sansevieria suffruticosa N. E. Br. (Asparagaceae Juss.)

The results of studing flower morphology, vascular anatomy and gynoecium micromorphology of Sansevieria suffruticosa N. E. Br. are presented. In this species, there is a flower tube of semiadnated tepals and stamens and also syncarpous gyno­ecium having septal nectaries. According to our data, the vascular system of this flower is comparable with the vascular system of the flower having free tepals, stamens and carpels because all of them have independent innervation. The carpel trace consists of dorsal bundle and a median ventral bundle which bifurcated in two ventral bundles, fused with dorsal bundle in the ovary roof. The ovule is innervated from the median ventral bundle. The gynoecium of Sansevieria suffruticosa is hemisyncarpous, but it has a short synascidiate zone which is typical for the eusyncarpous gynoecium. The hemisynascidiate (fertile), hemisymplicate (higher part of the ovary) and asymplicate (style and stigma) zones are located above a synascidiate zone. The septal nectary in the ovary corresponds to the inner non-labyrinthine distinct type, and in the ovary roof – to the outer labyrinthine type. The high base and roof of the ovary, and also long septal nectary are characteristic features of the inner gynoecium structure of this species. Keywords : Sansevieria , flower morphology, vascular anatomy, gynoecium, septal nectaries.

A guide to the gynoecium

Understanding flower structure is important from many perspectives such as keying out plants, understanding fruit structure, investigating pollinator biology and in plant breeding. Probably the most complicated parts of a flower are the female components (the gynoecium). Unfortunately, in many parts of the world, it is not possible for much of the year to provide students with the diversity of flowers needed to show an adequate range of gynoecium structure. To overcome these limitations I have developed a web-based tutorial that uses quality photographic images of fresh material of about 50 species to show a wide diversity of gynoecium structure. The tutorial is supported with an interactive test based on 12 species not used in the tutorial. Combined, the tutorial and test allow students to quickly become familiar with a full range of gynoecium structure. The online availability allows students to come better prepared for their introductory flower structure practicals and to review these materials at a later date.

Comparative floral morphology and anatomy of Anacardiaceae and Burseraceae (Sapindales), with a special focus on gynoecium structure and evolution

Anacardiaceae and Burseraceae are traditionally distinguished by the number of ovules (1 vs. 2) per locule and the direction of ovule curvature (syntropous vs. antitropous). Recent molecular phylogenetic studies have shown that these families are sister groups in Sapindales after having been separated in different orders for a long time. We present a comparative morphological study of the flower structure in both families. The major clades, usually supported in molecular phylogenetic analyses, are well supported by floral structure. In Anacardiaceae, there is a tendency to gynoecium reduction to a single fertile carpel (particularly in Anacardioideae). The single ovule has a long and unusually differentiated funicle, which connects with the stylar pollen tube transmitting tract in all representatives studied. In Anacardiaceae–Spondiadoideae, there is a tendency to form an extensive synascidiate zone, with a massive remnant of the floral apex in the centre; these features are also present in Beiselia (Burseraceae) and Kirkiaceae (sister to Anacardiaceae plus Burseraceae) and may represent a synapomorphy or apomorphic tendency for the three families. In core Burseraceae, gynoecium structure is much less diverse than in Anacardiaceae and has probably retained more plesiomorphies. Differences in proportions of parts of the ovules in Anacardiaceae and Burseraceae are linked with the different direction of ovule curvature. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159, 499–571.

Comparative Gynoecium Structure and Development in Calycanthaceae (Laurales)

Calycanthaceae, sister to all other Laurales, are the most distinct family of the order in their gynoecium structure (lateral placentation, presence of two ovules). Gynoecium structure was studied in a representative of each genus of Calycanthaceae and gynoecium development in a representative of each of the two subfamilies (Calycanthoideae and Idiospermoideae). Newly found shared features are postgenital coherence between the free carpels (extragynoecial compitum), abortion of the upper of the two ovules, and lobation of the outer integument. Differences in reproductive structures between the two subfamilies are reviewed. Newly found differences include carpel primordium shape, contribution of the outer integument to micropyle formation, epidermis differentiation of the rim of the integuments, and mode of formation of the compitum. Unexpectedly, at anthesis, ovary and ovules of Idiospermum are not larger than those of Calycanthoideae, despite the conspicuous difference in fruit. The identity of “staminod...

Flower structure and development in Tupidanthus calyptratus (Araliaceae): an extreme case of polymery among asterids

Flowers of Tupidanthus show an extreme case of floral polymery among asterids. Floral development and gynoecium structure have been examined. The floral meristem has a complex folded shape. The tiny calyx is initiated as a continuous ring primordium. The corolla is initiated as a lobed ring and develops into a calyptra. All stamen primordia appear simultaneously as a single whorl. The carpels, also in a single whorl, tend to alternate with the stamens. Some Schefflera species related to Tupidanthus are also studied. The flower of Tupidanthus is interpreted as a result of fasciation. Further investigation should determine whether mutation(s) in gene(s) of the CLAVATA family are responsible for the fasciation here. The significance of Tupidanthus for understanding spatial pattern formation in flowers of Araliaceae, and both functional and developmental constraints in angiosperm flowers with a single polymerous carpel whorl are discussed.

Cunoniaceae in the Cretaceous of Europe: Evidence from Fossil Flowers

Fossil flowers of the Cunoniaceae from Late Cretaceous sediments of southern Sweden are described in detail. The flowers are small, bisexual, actinomorphic, tetramerous with broadly attached valvate sepals; they have narrowly attached petals; eight stamens in two whorls; a massive, lobed nectary; a semi-inferior, syncarpous gynoecium with axile placentation; numerous ovules; separate styles; and peltate, probably secretory, trichomes. They share many features with extant representatives of both the Cunoniaceae and Anisophylleaceae. However, the gynoecium structure in particular indicates a closer relationship to the Cunoniaceae. The floral characters are not specific for any extant genus of the family and therefore a new genus and species, Platydiscus peltatus gen. et sp. nov., is formally described. This is the first record of cunoniaceous floral structures from the Northern Hemisphere and the oldest record of Cunoniaceae flowers worldwide.

Gynoecium diversity and systematics in basal monocots

Gynoecium and ovule structure was comparatively studied in representatives of the basal monocots, including Acorales (Acoraceae), Alismatales (Araceae, Alismataceae, Aponogetonaceae, Butomaceae, Hydrocharitaceae, JuncÍaginaceae, Limnocharitaceae, Potamogetonaceae, Scheuchzeriaceae, Tofieldiaceae), Dioscoreales (Dioscoreaceae, Taccaceae), and Triuridaceae as a family of uncertain position in monocots. In all taxa studied the carpels or gynoecia are closed at anthesis. This closure is attained in different ways: (1) by secretion without postgenital fusion (Araceae, Hydrocharitaceae); (2) by partly postgenitally fused periphery but with a completely unfused canal (Alismataceae, Aponogetonaceae, Butomaceae, Limnocharitaceae, Scheuchzeriaceae, Dioscoreaceae, Taccaceae); (3) by completely postgenitally fused periphery but with an unfused canal in the centre (Acoraceae, Tofieldiaceae); (4) by complete postgenital fusion and without an (unfused) canal (Juncaginaceae, Potamogetonaceae). In many Alismatales (but without Araceae) carpels have two lateral lobes. The stigmatic surface is restricted to the uppermost part of the ventral slit (if the carpel is plicate); it is never distinctly double-crested (Butomaceae?). Stigmas are commonly unicellular-papillate and secretory in most taxa. The locules are filled with a (often) mucilaginous secretion in a number of taxa. Superficial (probably intrusive) ethereal oil cells were found in the carpel wall of Acorus gramineus (as in Piperales!). Idioblasts in carpels are otherwise rare. A number of basal monocots has orthotropous ovules, which is perhaps the plesiomorphic condition in the group. The presence of almost tenuinucellar (pseudocrassinucellar) ovules is relatively common (Acoraceae, many Araceae, some Alismatales s.s.), whereas completely tenuinucellar ovules are rare and do not characterize larger groups. However, crassinucellar ovules occur in the largest number of families among the study group (basal Araceae, many Alismatales s.s.) The outer integument is always annular in orthotropous ovules. The inner integument is often lobed and it mostly forms the micropyle, whereas the outer integument is always unlobed. Gynoecium structure supports the isolated position of Acoraceae as sister to all other monocots. However, in an overall view, if compared with all other families, Acoraceae clearly shows the greatest similarities with Araceae.

Gynoecium diversity and systematics in basal monocots

Abstract Gynoecium and ovule structure was comparatively studied in representatives of the basal monocots, including Acorales (Acoraceae), Alismatales (Araceae, Alismataceae, Aponogetonaceae, Butomaceae, Hydrocharitaceae, JuncIaginaceae, Limnocharitaceae, Potamogetonaceae, Scheuchzeriaceae, Tofieldiaceae), Dioscoreales (Dioscoreaceae, Taccaceae), and Triuridaceae as a family of uncertain position in monocots. In all taxa studied the carpels or gynoecia are closed at anthesis. This closure is attained in different ways: (1) by secretion without postgenital fusion (Araceae, Hydrocharitaceae); (2) by partly postgenitally fused periphery but with a completely unfused canal (Alismataceae, Aponogetonaceae, Butomaceae, Limnocharitaceae, Scheuchzeriaceae, Dioscoreaceae, Taccaceae); (3) by completely postgenitally fused periphery but with an unfused canal in the centre (Acoraceae, Tofieldiaceae); (4) by complete postgenital fusion and without an (unfused) canal (Juncaginaceae, Potamogetonaceae). In many Alismatales (but without Araceae) carpels have two lateral lobes. The stigmatic surface is restricted to the uppermost part of the ventral slit (if the carpel is plicate); it is never distinctly double-crested (Butomaceae?). Stigmas are commonly unicellular-papillate and secretory in most taxa. The locules are filled with a (often) mucilaginous secretion in a number of taxa. Superficial (probably intrusive) ethereal oil cells were found in the carpel wall of Acorus gramineus (as in Piperales!). Idioblasts in carpels are otherwise rare. A number of basal monocots has orthotropous ovules, which is perhaps the plesiomorphic condition in the group. The presence of almost tenuinucellar (pseudocrassinucellar) ovules is relatively common (Acoraceae, many Araceae, some Alismatales s.s. ), whereas completely tenuinucellar ovules are rare and do not characterize larger groups. However, crassinucellar ovules occur in the largest number of families among the study group (basal Araceae, many Alismatales s.s. ) The outer integument is always annular in orthotropous ovules. The inner integument is often lobed and it mostly forms the micropyle, whereas the outer integument is always unlobed. Gynoecium structure supports the isolated position of Acoraceae as sister to all other monocots. However, in an overall view, if compared with all other families, Acoraceae clearly shows the greatest similarities with Araceae.

Gynoecium Structure and Evolution in Basal Angiosperms

A comparative study on gynoecium structure in basal angiosperms revealed new features relevant to systematics. In addition, based on the new phylogenetic framework on basal angiosperms gained by molecular systematics, an evolutionary evaluation of gynoecial character states has become possible. Angiospermy patterns are diverse and are here tentatively grouped into four types according to the mode of carpel closure: (1) by secretion, without postgenital fusion; (2) with a continuous secretory canal but partial postgenital fusion at the periphery; (3) with an incomplete secretory canal and complete postgenital fusion at the periphery; (4) by complete postgenital fusion. In most basal angiosperms the carpels are free, and the inner space of each carpel is occluded from the outside by secretion and not by postgenital fusion (angiospermy type 1). Correlated with this feature is a pronounced ascidiate shape of the carpels. The carpels tend to be shortly stipitate. In relatively many basal clades with pluricarpellate gynoecia an extragynoecial compitum is present. Stigmas are secretory and tend to have pluricellular papillae. Ethereal oil cells at the surface of the carpels (probably intrusive) are present in Illiciales (including Trimeniaceae), Chloranthaceae, Piperales s.l. (including Aristolochiales), and Acoraceae. Ovule structure is diverse. Ovules are predominantly anatropous. Although Amborellaceae, the phylogenetically earliest branching extant angiosperm, has orthotropous ovules, it is argued that the anatropous condition is basal in angiosperms. Crassinucellar ovules are predominant in basal angiosperms. However, there are apomorphic tendencies toward ovule reduction: weakly crassinucellar (Nymphaeales, some Alismatales), pseudocrassinucellar (Acorales, some Alismatales, some Ranunculales), and tenuinucellar (some Piperales s.l., including Aristolochiales, Rafflesiales, some Alismatales). Lobation of the inner integument and semiannular shape of the outer integument are variously present or absent. They appear to be due to architectural constraints in ovule development and not to constitute archaic features. In contrast to anatropous ovules, orthotropous ovules often have an outer integument that is thinner than the inner one. This may be due to a function of the outer integument in ovule curvature as indicated by molecular developmental genetics. On the other hand, the presence of an outer integument that is thinner than the inner one characterizes the majority of the Piperales s.l. (including Aristolochiales), irrespective of presence or absence of ovule curvature.

Gynoecium diversity and systematics of the basal eudicots

Gynoecium and ovule structure was compared in representatives of the basal eudicots, including Ranunculales (Berberidaceae, Circaeasteraceae, Eupteleaceae, Lardizabalaceae, Menispermaceae, Papaveraceae, Ranunculaceae), Proteales (Nelumbonaceae, Platanaceae, Proteaceae), some families of the former «lower» hamamelids that have been moved to Saxifragales (Altingiaceae, Cercidiphyllaceae, Daphniphyllaceae, Hamamelidaceae), and some families of uncertain position (Gunneraceae, Myrothamnaceae, Buxaceae, Sabiaceae, Trochodendraceae). In all representatives studied, the carpels (or syncarpous gynoecia) are closed at anthesis. This closure is attained in different ways: (1) by secretion without postgenital fusion (Berberidaceae, Papaveraceae, Nelumbonaceae, probably Circaeaster); (2) by a combination of postgenital fusion and secretion; (2a) with a complete secretory canal and partly postgenitally fused periphery (Lardizabalaceae, Menispermaceae, some Ranunculaceae, Sabiaceae); (2b) with an incomplete secretory canal and completely fused periphery (Trochodendron); (3) by complete postgenital fusion without a secretory canal (most Ranunculaceae, Eupteleaceae, Platanaceae, Proteaceae, all families of Saxifragales and incertae sedis studied here). Stigmas are double-crested and decurrent in most of the non-ranunculalian taxa; unicellular-papillate in most taxa, but with multicellular protuberances in Daphniphyllaceae and Hamamelidaceae. Carpels predominantly have three vascular bundles, but five in Proteales (without Nelumbonaceae), Myrothamnaceae and Trochodendraceae. The latter two also share «oil» cells in the carpels. Stomata on the outer carpel surface are present in the majority of Ranunculales and Proteales, but tend to be lacking in the saxifragalian families. In basal eudicots, especially in the non-ranunculalian families there is a trend to form more than one ovule per carpel but to develop only one seed, likewise there is a trend to have immature ovules at anthesis. Ovule number per carpel is predominantly one or two. Proteales (without Nelumbonales) mainly have orthotropous ovules, the other groups have anatropous (or hemitropous or campylotropous) ovules. The outer integument is annular in the groups with orthotropous or hemitropous ovules, and also in a number of saxifragalian families with anatropous ovules. In Proteales the integuments are predominantly lobed but there is no distinct pattern in this feature among the other groups. Among Ranunculales two pairs of families (Lardizabalaceae/Menispermaceae and Berberidaceae/Papaveraceae) due to similarities in gynoecium structure can be recognized, which are not apparent in molecular analyses. The close relationship of Platanaceae and Proteaceae is supported by gynoecium structure but gynoecial features do not support their affinity to Nelumbonaceae. The alliance of Daphniphyllaceae with Hamamelidaceae s.l. is also supported.