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Drought, salinization and heavy metal pollution of soils are main stress factors with an increasing impact on the deterioration of soil quality, yield and crop quality. Seed priming shows good results in improving seed germination, seedling growth and plant development. Proline (Pro) and metyl jasmonate (MeJA) show stimulating activity and help plants overcome stress. The study investigated the effect of Pro, MeJA and hydropriming on seeds sown on water agar supplemented with different concentrations of heavy metals (Cd, Pb, Zn) (HM), NaCl or Polyethylene glycol 6000 (PEG 6000). Chelidonium majus is a medicinal species which is grown as a crop in some parts of Europe. It is an ingredient in some remedies and is becoming an increasingly popular object of research regarding its biological activities. The low concentrations of all heavy metals applied increased the germination of all variants of seeds – control, hydroprimed and those which were Pro and MeJA primed. Seed priming with Pro and MeJA promoted high germination percentage of seeds germinated on water agar with NaCl. PEG 6000 at its higher concentration (5%) slightly increased the seed germination of all variants. The growth of roots and hypocotyls was inhibited by HM and NaCl. However, PEG 6000 slightly influenced their growth.

Coreanomecon is a monotypic and endemic genus in Korea, distributed mainly in the southern regions. Coreanomecon is morphologically similar to Hylomecon by producing red latex, easily distinguished from Chelidonium, which produces yellow latex. Coreanomecon were merged into Hylomecon or Chelidonium depending on the authors. To understand the phylogenetic relationship of Coreanomecon, DNA sequences of chloroplast rbcL and matK and nuclear Internal Transcribed Spacer (ITS) regions were determined from the species of Papaveroideae (Papaveraceae) in Korea and analyzed with the Maximum Parsimony and Bayesian methods. Phylogenetic analyses of Papaveroideae suggest that Coreanomecon is sister to the clade of Chelidonium and Stylophorum in the ITS data and that it is sister to Hylomecon in the chloroplast (cpDNA) data. A constraining analysis using the Shimodaira-Hasegawa test (S-H test) suggested that the ITS data do not reject the sister relationship of Coreanomecon and Hylomecon. The S-H test also suggested that the cpDNA data is compatible with the placement of Coreanomecon as a sister to the clade of Chelidonium and Stylophorum. Although the conflicting phylogenetic results may stem from insufficient phylogenetic signals, they may also be associated with hybridization between Hylomecon and an ancestor of Stylophorum and Chelidonium. The results of this study suggest that Coreanomecon is a distinct lineage as an endemic genus, supporting the morphological data.

The complete plastome of Sanguinaria canadensis was sequenced, which is 160,117 bp in length, consisting of 87699 bp large single-copy (LSC), a 19,136 bp small single copy (SSC), and a pair of 26,638 bp inverted repeat (IR) regions. The S. canadensis plastome encodes 137 annotated genes including 37 tRNA genes, 8 rRNA genes, and 89 protein coding genes. Phylogenetic analysis strongly supported that it is a member of Chelidonieae.

Macleaya cordata (Willd.) is a perennial herb with significant antimicrobial and anti-inflammatory activities. In this study, we report the complete chloroplast genome sequence of M. cordata. The t...

— To explore the taxonomic status, phylogenetic patterns, and character evolution of various groups within Papaveraceae s. l. (Pteridophylloideae, Papaveroideae, Hypecoideae, and Fumarioideae), two data sets were created and analyzed using Bayesian inference, maximum likelihood, and parsimony methods: 1) a Ranunculales data set for 82 genera representing all of the major families within the order using three chloroplast DNA regions (atpB, rbcL, and matK) and nuclear 26S ribosomal DNA, and 2) a Papaveraceae s. l. data set for an expanded sampling of the family (27 genera, 40 species) consisting of chloroplast sequences only. Branch support values on the resulting trees derived from the combined data for both the Ranunculales and Papaveraceae s. l. data sets were comparable among the various ranunculalean families and supported subfamilial recognition of Pteridophylloideae, Papaveroideae, Hypecoideae, and Fumarioideae. Within Papaveraceae s. l., most analyses of both data sets resolved the position ...

Chemical investigation of the aerial parts of Meconopsis horridula led to the isolation and structural identification of nine flavonoids, two alkaloids and five terpenoids. Their structures were determined by comparison of 1H and 13C NMR data with those reported in literature. The discovery of alkaloid cavidilinine (11) and two pentacyclic triterpenoids (15 and 16) from M. horridula provides evidence to support the view that Fumarioideae and Papaveroideae should be combined into a single family Papaveraceae.

International audience; Premise of the Study: Zygomorphy has evolved multiple times in angiosperms. Near-actinomorphy is the ancestral state in the early diverging eudicot family Papaveraceae. Zygomorphy evolved once in the subfamily Fumarioideae from a disymmetric state. Unusual within angiosperms, zygomorphy takes place along the transverse plane of the flower. Methods: We investigated floral development to understand the developmental bases of the evolution of floral symmetry in Papaveraceae. We then assessed the expression of candidate genes for the key developmental events responsible for the shift from disymmetry to transverse zygomorphy, namely CRABSCLAW for nectary formation (PAPCRC), SHOOTMERISTEMLESS (PAPSTL) for spur formation, and CYCLOIDEA (PAPCYL) for growth control. Key Results: We found that an early disymmetric groundplan is common to all species studied, and that actinomorphy was acquired after sepal initiation in Papaveroideae. The shift from disymmetry to zygomorphy in Fumarioideae was associated with early asymmetric growth of stamen filaments, followed by asymmetric development of nectary outgrowth and spur along the transverse plane. Patterns of PAPSTL expression could not be clearly related to spur formation. PAPCRC and PAPCYL genes were expressed in the nectary outgrowths, with a pattern of expression correlated with asymmetric nectary development in the zygomorphic species. Additionally, PAPCYL genes were found asymmetrically expressed along the transverse plane in the basal region of outer petals in the zygomorphic species. Conclusion: Genes of PAPCRC and PAPCYL families could be direct or indirect targets of the initial transversally asymmetric cue responsible for the shift from disymmetry to zygomorphy in Fumarioideae.

Since the nineteenth century, the western European endemic Meconopsis cambrica has been regarded as the only European representative of the genus Meconopsis Vig. This genus, which is otherwise restricted to the Himalayas, differs from Papaver in having a style rather than a stigmatic disc. A phylogenetic reconstruction using 65 internal transcribed spacer sequences of 62 taxa of Old World Papaveroideae and three outgroup taxa shows that M. cambrica is not the closest relative of the remainder of Meconopsis but rather the closest sampled relative of Papaver s.str. This is consistent with morphological evidence which suggests that the style evolved independently in M. cambrica from a Papaver-like ancestor. We suggest that Meconopsis cambrica should revert to its former name, Papaver cambricum L. We also suggest that: (1) the Arctic-montane species of Papaver sect. Meconella need to be placed in their own genus; (2) Papaver sect. Argemonidium (including P. argemone and P. hybridum) should be included...

International audience; Background and Aims Fumarioideae (20 genera, 593 species) is a clade of Papaveraceae (Ranunculales) characterized by flowers that are either disymmetric (i.e. two perpendicular planes of bilateral symmetry) or zygomorphic (i.e. one plane of bilateral symmetry). In contrast, the other subfamily of Papaveraceae, Papaveroideae (23 genera, 230 species), has actinomorphic flowers (i.e. more than two planes of symmetry). Understanding of the evolution of floral symmetry in this clade has so far been limited by the lack of a reliable phylogenetic framework. Pteridophyllum (one species) shares similarities with Fumarioideae but has actinomorphic flowers, and the relationships among Pteridophyllum, Papaveroideae and Fumarioideae have remained unclear. This study reassesses the evolution of floral symmetry in Papaveraceae based on new molecular phylogenetic analyses of the family. Methods : Maximum likelihood, Bayesian and maximum parsimony phylogenetic analyses of Papaveraceae were conducted using six plastid markers and one nuclear marker, sampling Pteridophyllum, 18 (90 %) genera and 73 species of Fumarioideae, 11 (48 %) genera and 11 species of Papaveroideae, and a wide selection of outgroup taxa. Floral characters recorded from the literature were then optimized onto phylogenetic trees to reconstruct ancestral states using parsimony, maximum likelihood and reversible-jump Bayesian approaches. Key Results : Pteridophyllum is not nested in Fumarioideae. Fumarioideae are monophyletic and Hypecoum (18 species) is the sister group of the remaining genera. Relationships within the core Fumarioideae are well resolved and supported. Dactylicapnos and all zygomorphic genera form a well-supported clade nested among disymmetric taxa. Conclusions : Disymmetry of the corolla is a synapomorphy of Fumarioideae and is strongly correlated with changes in the androecium and differentiation of middle and inner tepal shape (basal spurs on middle tepals). Zygomorphy subsequently evolved from disymmetry either once (with a reversal in Dactylicapnos) or twice (Capnoides, other zygomorphic Fumarioideae) and appears to be correlated with the loss of one nectar spur.

To determine the relationship among the Papaveroideae, a data set, originally gathered for classification of the Papaveroideae using DNA restriction sites, was subjected to an multidimensional scaling (MDS) analysis, leading to a more appropriate taxonomy of the Papavereae tribe (especially in Papaver, Stylomecon, and Roemeria genera). The spatial configuration of Papavereae was developed on the basis of absolute distances of taxa in relation to Papaver alpinum and made possible the 3D projection of the tribe. The configuration consisted of two different sets and provided a divergent rank image of taxa from the cladistic assumption. In one of these sets, taxa of Roemeria and the Papaver section Argemonidium, were not mixed, but formed two distinct groups in view of the genera. Papaver hybridum was the closest species to Roemeria, but Papaver pavoninum was the farthest and most distant taxon. In case of annual poppy species of the other set, the Papaver somniferum and Papaver macrostomum were not ...

Evolutionary relationships withinArgemone (Papaveraceae) were inferred from nucleotide variation in nuclear ribosomal DNA. A complete representation of the genus was achieved by using herbarium material to a large extent (74%). Four distinct clades can be recognized based on the molecular results. The support for the different clades varies greatly due to a very uneven distribution of characters. Although some clades are largely unresolved, some unexpected relationships, for example the sister group relationship of the relict speciesA. subintegrifolia to the rest ofArgemone, were found. The evolution of morphological and alkaloid characters against the background of the obtained phylogeny is discussed. Whereas the morphological characters provide some support for the phylogeny found, the alkaloid characters turned out not to contain much phylogenetic information. Past difficulties in recognizing phylogenetic relationships withinArgemone result from largely continuous morphological variation and from parallelisms correlated with and probably caused by polyploidization.

Summary This study presents an analysis of leaf development in 13 taxa of Papaveraceae subfamily Papaveroideae. The emphasis is on the organogenetic phase of leaf growth, during which leaflet and serrature primordia are initiated. The patterns of organogenetic activity are visualized by scanning electron microscopy and schematic drawings, and are related to regional morphogenetic and histogenetic processes, particularly thiqkening growth, surface growth, margin dilatation, and differential histogenesis. In Romneya coulteri, leaflet initiation, leaflet segmentation, and the course of higher-order segmentation proceed acropetally. In contrast, the respective patterns are basipetal in Argemone. The entire leaves of Platystemon lack segmentation. Within the closely related genera Papaver, Roemeria and Meconopsis, acropetal (Roemeria refracta) and divergent (Papaver atlanticum, P. pilosum) modes of leaflet initiation are connected by intermediate patterns (Papaver pavoninum, P. armeniacum, P. dubium,P. somniferum and Meconopsis cambrica). These are characterized by a divergent wave of higher-order segmentation following acropetal or simultaneous-acropetal leaflet formation. The construction of a morphocline indicates that heterochronic changes account for leaf diversification in this group. Meconopsis betonicifolia exhibits simple basipetal segmentation. Comparative observations of leaf morphogenesis in the above taxa indicate that differential marginal dilatation and histogenesis independently contribute to the determination of dissection patterns, and that leaf shape diversity arises from the potential to generate various combinations of regional growth processes.

During the present course, population-based meiotic studies were carried out on five species of subfamily Papaveroideae from selected localities of Kashmir and Himachal Pradesh in the Western Himalayas (India). Varied intraspecific chromosome counts were reported for the first time in Argemone mexicana and Meconopsis latifolia, both existing on 2n = 2x = 14. The x = 7, confirmed for the first time from the newly found diploid cytotype, is suggested to be the primary chromosomal basic number for the Meconopsis. Furthermore, meiotic course was noted to be normal in Argemone ochroleuca, it varied from normal to abnormal in the populations of A. mexicana and Papaver dubium whereas it was invariably found to be abnormal in all the populations of Meconopsis aculeata and M. latifolia. These anomalous taxa were marked with meiotic abnormalities in the form of cytomixis, chromosomal stickiness, unoriented bivalents, formation of laggards and bridges resulting in abnormal microsporogenesis, and production of heterogeneous-sized fertile pollen grains along with reduced pollen fertility.

PREMISE OF THE STUDY: The correct assessment of homology is an important prerequisite for reconstructing phylogenetic relationships and character evolution. Old World Papaveroideae (Papaver, Meconopsis, Roemeria, Stylomecon) show substantial diversity in gynoecium and capsule morphology. In particular, capsules can have distinct styles (Meconopsis p.p., Stylomecon) or a sessile stigmatic disc (Papaver). Molecular phylogenetic analyses of Old World Papaveroideae had shown that neither taxa with styles nor those with stigmatic discs represent monophyletic lineages. We here investigate whether either styles or stigmatic discs have arisen repeatedly during the diversification of Old World Papaveroideae. METHODS: We investigated gynoecium ontogeny in Papaver rhoeas, P. californicum, Meconopsis cambrica, and Stylomecon heterophylla by scanning electron microscopy for the first time. Our observations were interpreted on the background of a well-resolved molecular phylogeny of the taxa investigated. KEY RESULTS: Papaver rhoeas and P. californicum share the presence of a developmentally complex garland-like stage in gynoecium ontogeny. The styles of M. cambrica and S. heterophylla result from growth in a ring-like zone beneath the carpel tips. This zone is also present in Papaver. In M. cambrica, traces of a garland-like stage can be seen. Style formation and stigma formation begin more or less simultaneously in M. cambrica, but style formation clearly precedes stigma formation in S. heterophylla. CONCLUSIONS: The styles present in M. cambrica and S. heterophylla are considered to have arisen in parallel from ancestors with a stigmatic disc. We speculate that style formation may have been a means to reduce selfing.

The RFLP-analysis of PCR amplified cpDNA fragments of 42 representatives of Papaveraceae subfam. Papaveroideae resulted in six most parsimonious cladograms. The subfamily can be divided into a New World group (Arctomecon, Argentone, Canbya, Romneya and Platystemon) and an Old World group (Meconopsis, Papaver s.l. and Roemeria) containing Stylomecon heterophylla and Papaver californicum as New World taxa. In the Old World group neither Meconopsis nor Papaver are monophyletic. Whereas Meconopsis consists of three clades, Papaver comprises five clades, with Roemeria as sister group to P. sect. Argemonidium and Stylomecon as sister group to P. californicum. Various lines of evidence suggest that Meconopsis should be interpreted as a paraphyletic base group of Old World Papaveroideae which gave rise to a polyphyletic Papaver.

On the background of the phylogeny of the Papaveraceae s.l., the evolution of the androecium and gynoecium, the geographical distribution, and the ecological differentiation of the family is discussed. An effort is made to homologize the diverse androecia of Pteridophyllum, Hypecoum, Fumariaceae and Papaveraceae s.str., and it is concluded that both the polyandrous androecium as found in the Papaveraceae s.str. and the polycarpellate gynoecium as found in Papaveraceae s. str. subfam. Papaveroideae incl. Platystemonoideae are secondary and derived from whorled oligomerous androecia and bicarpellate gynoecia, respectively. The comparative consideration of the geographical distribution, ecology and diversity of various monophyletic units within the family suggests (1) that forest floor habitats might be ancestral in the family, (2) that in parts of the family generic diversification preceded the break-up of the northern continents, and (3) that aridification of the environment was a major stimulus for both diversification and morphological divergence in the family.

A morphological-based cladistic analysis was conducted on 116 ingroup taxa and 5 outgroups coded for 109 characters and 192 apomorphic character states. Ranunculid genera are better organized as the two orders Lardizabalales and Ranunculales, rather than Ranunculales and Papaverales of contemporary systems. Within Lardizabalales, the genus Decaisnea is segregated as a new family, in contrast to Sargentodoxa, which is maintained within Lardizabalalceae. The Berberidaceae and Menispermaceae are early branches of Ranunculales, which is primitively woody. Hydrastidaceae (including Glaucidium), Papaveraceae, and Ranunculaceae form a herbaceous clade of derived Ranunculales. The Papaveraceae subfamilies Platystemonoideae, Papaveroideae, and Eschscholzioideae are early branches, whereas Pteridophylloideae, Hypoecoideae, and Fumarioideae are derived. The Chelidonioideae are transitional and paraphyletic and therefore, a new subfamily is recognized for Glaucium and Dicranostigma. The Ranunculaceae subfamily lsopyroideae is a first branch, and Helleboroideae and Ranunculoideae are sister graups. Within lsopyroideae, the Aquilegieae is a first branch, and lsopyreae and Coptidae (including Dichocarpum and Thalictrum) are sister groups. Within Helleboroideae, the Caltheae is a first branch, and Helleboreae (including Nigella) and Cimicifugeae (including Aconitum, Beesia, and Delphinium) are sister groups. Within Ranunculoideae, the Adonideae is a first branch, and Anemoneae and Ranunculeae are sister groups. Circaeaster and Kingdonia are sister groups and positioned as a first branch of the Anemoneae, and it is at least 7 steps less parsimonious to remove both from Ranunculaceae, and 14 steps longer to remove only Circaeaster. It is 10 steps less parsimonious to consider a Ranunculaceae ancestry for Paeonia, which is positioned in Dilleniales. The “ranunculidae” are paraphyletic because Ranunculales are closer to Dilleniales than to Lardizabalales.

Phylogenetic relationships of a group of North American desert genera of the Papaveraceae subfam. Papaveroideae and Platystemonoideae were investigated with an RFLP analysis of three PCR-amplified chloroplast genome regions. In agreement with earlier results it was found that subfam. Platystemonoideae is nested within subfam. Papaveroideae. The group under study is characterized by a large number of generic autapomorphies but only few informative synapomorphies. This is interpreted as strong evidence for a rapid radiation event caused by major climatic changes in the past. Poor phylogenetic resolution seems to reflect biological reality and not to be an experimental artifact. There is also evidence for accelerated morphological evolution under arid climatic conditions.

Phylogenetic analyses of various combinations of atpB and rbcL sequence data, trnK restriction sites, and morphological characters of taxa representing the Papaveraceae s.l. (Pteridophyllaceae, Fumari- aceae subfamilies Fumarioideae and Hypecoideae, Papaveraceae s. str. subfamilies Eschscholzioideae, Chelidonioideae, Papaveroideae, Platystemonoideae) are presented. Several measures of data set incongru- ency indicate considerable congruence between data sets. In order to correctly identify the root of the family, combined atpB and rbcL sequence data for the Papaveraceae s.l. and appropriate outgroups (Asarum, Illicium, Platanus, Euptelea, and representatives of the Ranunculaceae and Berberidaceae) were analyzed. Regardless of the outgroup used, all topologies obtained suggest that Pteridophyllum (Pteridophyllaceae) is sister group to Fumariaceae and Papaveraceae s. str. Data sets of the ingroup with Pteridophyllum used as the outgroup were analyzed separately and together, with results from the combined data sets showing increased bootstrap support. Contrary to recent claims, Papaveraceae s. str. remain monophyletic after the exclusion of Pteridophyl- lum and Fumariaceae. In the Papaveraceae s. str., Eschscholzioideae are sister group to Papaveroideae (including Platystemonoideae) and Chelidonioideae (including Glaucium/Dicranostigma). The evolution of floral morphology, geographical distribution, and ecology of Papaveraceae s.l. are briefly discussed.

Basierend auf vergleichend-morphologischen Untersuchungen an 20 Papaveraceae-Gattungen wird eine Beschreibung der Fruchtmorphologie (incl. Anatomie) fur jede Gattung gegeben. Bei 4 Gattungen konnte lediglich auf Angaben aus der Literatur zuruckgegriffen werden. Fur die 4 Triben sind bestimmte Kombinationen von Differenzierungsmerkmalen charakteristisch. Besondere Beachtung fanden dabei die Struktur der auseren und inneren Perikarp-Epidermis, die Chlorenchym-Verteilung, der Verlauf der Milchrohren, die Gynoeceumsvaskulatur und die Dehiszenzform. Die homokarpellaten Platystemoneae (Unterfamilie Platystemonoideae) sind von den heterokarpellaten Papaveroideae klar abgegrenzt. Die Chelidonieae sind u. a. nach den Dehiszenzformen die plastischste Tribus, hier finden sich zahlreiche ursprungliche Merkmalsauspragungen. Die sich explosiv akropetal offnenden Kapseln der Eschscholzieae sind anatomisch relativ einheitlich. In den Papavereae treten nur unvollstandig basipetal dehiszierende Fruchte auf, die durch fehlende Valvendorsalnerven und im Phloem verlaufende Milchrohren charakterisiert sind. Bestimmte Strukturen weisen auf nahere Bezichungen zwischen Einzelgattungen hin, jedoch rechtfertigt dies keine taxonomische Neugliederung der Familie. Die Hypecoaceae stehen den Chelidonieae (Glaucium, Dicranostigma) und den Eschscholzieae nahe. — Ein phylogenetisches Schema zeigt die karpomorphologischen Beziehungen in den Papaveraceae. Based on comparative morphological capsule studies (incl. anatomy) of 20 papaveraceous genera a description of the fruit structure is given for each genus. 4 genera could be described by literature analysis only. Each tribe is characterized by a defined set of differentiation characters. The following features have been examined in particular: structure of outer and inner pericarp epidermis, localization of chlorenchyma, course of laticifers, vascularization of the gynoceum, and mode of dehiscence. The homocarpellate Platystemoneae (subfam. Platystemonoideae) are clearly separated from the heterocarpellate Papaveroideae. The Chelidonieae are the most plastic tribe concerning dehiscence and other features, they have retained many primitive character states. The capsules of the Eschscholzieae opening violently acropetally are relatively uniform in anatomical respect. In the Papavereae there are incompletely basipetally dehiscing fruits which are characterized by lacking dorsal valve traces and laticifers restricted to the pholem. Special structures indicate closer relations between individual genera but do not demand a new arrangement of the taxa. The Hypecoaceae show comparatively strong affinities to Chelidonieae (Glaucium, Dicranostigma) and Eschscholzieae. — A phylogenetic diagram illustrates the carpomorphological relationships in the Papaveraceae.

Die Struktur der parakarpen Gynoeceen in der Unterfamilie Papaveroideae der Familie Papaveraceae Juss. wird seit uber 150 Jahren unterschiedlich interpretiert. Neben der Auffassung, das das Gynoeceum so viele Karpelle wie Valven besitzt, existiert die Vorstellung, das unterschiedliche Karpellformen (fertile “solide” bzw. “kontrahierte” Karpelle und sterile “Valven-” bzw. “expandierte” Karpelle) am Aufbau beteiligt sind. Zur Klarung dieser Frage wurden die Gynoeceen zahlreicher Gattungen morphologisch untersucht, wobei der Vaskulatur der Plazentarregionen besondere Aufmerksamkeit geschenkt wurde. Verschiedene teratologische Gynoeceen wurden ebenfalls unter diesem Aspekt ausgewertet. Die Ergebnisse lassen den Schlus zu, das die Gynoeceen in der Unterfamilie Papaveroideae aus Karpellen zweier Formen zusammengesetzt sind. Valvenkarpelle alternieren mit mehr oder weniger stark kontrahierten Karpellen, die Hauptbestandteil der Plazentarregionen sind. Beide Karpellformen sind jedoch fertil. In der Unterfamilie Platystemonoideae ERNST tritt lediglich eine Karpellform auf, das “klassische” Karpell. Es wird eine Ubersicht uber die Herausbildung und die Evolutionstrends des Gynoeceums in der Familie gegeben. The structure of the paracarpous gynoecea in the subfamily Papaveroideae of Papaveraceae Juss. has been interpreted differently for more than 150 years. One conception is that there are as many carpels as valves in the gynoeceum. Another assumption postulates that the gynoeceum is composed of different carpel forms (fertile “solid” or “contracted” carpels and sterile “valve” or “expanded” carpels). To enlighten that problem the gynoecea of selected genera had been studied morphologically. Particular stress had been laid on the vascularization of the placental regions. Several teratological gynoecea had also been analysed regarding that aspect. The results allow the conclusion that there are really two different carpel forms being integrated in the papaveroideous gynoeceum. Valve carpels alternate with more or less contracted carpels that are the main constituents of the placental regions. However, both carpel forms are fertile. Only one carpel form, the “classical” carpel, occurs in the subfamily Platystemonoideae ERNST. A schematic representation of the origin and the evolutionary trends of the papaveraceous gynoeceum is given.

1) Many meiotic chromosome numbers in the Papaveraceae have been counted.2) Pteridophyllum is probably descended from a very remote ancestor of Chelidonioideae and should be put in the new family Pteriodophyllaceae.3) In the Papaveraceae there are two series, one of which, the Corydalo-Hypecoideae, having the basic chromosome number 4 and the other, Chelidonioideae the basic number 3. These two series are independent, but finally unite in Papaveroideae (e.g. Roemeria).There are two kinds of Roemeria: one with the basic number 3 and the other with 3 and 4, thus showing that Roemeria is karyologically a heterogenous genus.4) Glaucium which has been classified by Fedde with Papavereae, should be grouped with Chelidoneae (the writer's Chelidonioideae) as it is quite different karyologically from other genera in Papavereae.5) We conclude from the present studies that the following genera have been produced by crossing between plants of Corydaloideae and Chelidonioideae, but their crossing occurred early in the history of development of the family, and we are at present not in a position to trace the exact chromosomal history.*Hunnemannia 7=(4+3)………Corydaloideae (Corydalis, Dicentra or allied species)×EschscholtziaBocconia Macleaya 10=(4+3+3)……Corydaloideae (Corydalis, Dicentra or allied spedies)×ChelidoniumFumaria (Fumarioideae) 7=(4+3)………Corydalis or allied species×GlauciumPapaveroideae 7=(4+3) 11=(4+4+3)……Corydaloideae×Chelidonioideae6) Papaver can be said to be the most highly developed genus among the Papaveraceae, both for anatomical and karyological reasons.7) Fedde is of the opinion that Fumarioideae (Fumarioideae and Corydaloideae of the present scheme) is the youngest in this family, but as stated before, Fumaria, having a different number of chromosomes from other genera in Fumarioideae (the writer's Corydaloideae), seems to be derived from a genetic connection of Corydalis and Glaucium, as Fumaria and Glaucium have fumaric acid in common.The writer, therefore, believes that Fumaria is situated between Roemeria and Corydalis, having branched out from the latter. Other genera of Fumarioideae (the writer's Corydaloideae) also should be placed before the Papavereae (the writer's Papaveroideae) because of their having the basic number 4.8) Argemone, although it has fewer species as compared with Papaver, is the most advanced genus in America, while Papaver has become the most advanced genus in Europe and Asia Minor, having about 90 species.In conclusion I desire to express my best thanks to Hon. Prof. Fujii and Prof. Nakai, the director of the Botanical Institute of the Tokyo Imp. University for much valuable advice and the great interest they have shown in my work, and also to express my sincere appreciation of the help and encouragement I have received from Prof. Sinoto and Dr. Yasui during the course of the present work.

Publisher Summary This chapter provides an overview of the chemotaxonomy of two families: Papaveraceae and Fumariaceae. Both are very rich in specific alkaloids. The family Papaveraceae Juss. is subdivided into the subfamilies Hypecoideae Prantl and Kundig and Papaveroideae A. Br. It embraces 23 genera and ∼430 species. The Fumariaceae DC. include 7 genera with ∼350 species. The alkaloids of the Papaveraceae and Fumariaceae can be subdivided into several constitutional types—that is, simple isoquinolines, benzylisoquinolines, pavines, isopavines, cularines, proaporphines, aporphines, promorphinanes, morphinanes, protoberberines, retroprotoberberines, secoberbines, benzophenanthridines, protopines, phthalideisoquinolines, secophthalideisoquinolines, indenobenzazepines, spirobenzylisoquinolines, and rhoeadines. The most widespread alkaloids of the Papaveraceae Juss. are protoberberines, benzophenanthridines, and protopines, which, with a few exceptions, are present in all genera. The Papaveraceae and Fumariaceae DC. are closely related families. Their relationship and their independence are well demonstrated by the external structure of the flowers, the various anatomical characteristics of their different organs, and alkaloid content.

The genus Macleaya belongs to the family Papaveraceae Juss. (Jussieu 1789), subfamily Papaveroideae, tribe Chelidonieae Reichb. (Reichenbach 1837). As early as 1826, the genus Macleaya was separated from the genus Bocconia L. by Robert Brown. It includes two closely related species Macleaya cordata (Willd.) R. Br. (= Bocconia cordata Willd.) native to Central to Eastern China and Central Japan, and Macleaya microcarpa (Maxim.) Fedde (= Bocconia microcarpa Maxim.) which originated from the regions of Central China, Kansu and North Shensi (Fedde 1936). M cordata is an ornamental plant up to 3 m high and carrying splendid foliage. In 1795 it was introduced by G. Staunton into Middle European gardens. Common names for M cordata are Federmohn (German), plume poppy or tree celandine (English), takenigusa or champagiku (Japanese), for M microcarpa makleya melkoplodnaya (Russian).