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Most parsimonious course of evolution of perianth phyllotaxis on a tree with mesangiosperm relationships based on combined molecular and morphological data (D&E tree; Doyle & Endress, 2000; Endress & Doyle, 2009). Nymph = Nymphaeales, Aust = Austrobaileyales, Chlor = Chloranthaceae, Piper = Piperales, Ca = Canellales, Magnol = Magnoliales, OM = point of attachment of other monocots, OE = point of attachment of other eudicots. Modified from Endress & Doyle (2009).
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New morphological and phylogenetic data prompt us to present an updated review of floral morphology and its evolution in the basal ANITA grade of living angiosperms, Chloranthaceae, and Ceratophyllum. Floral phyllotaxis is complex whorled in Nymphaeales and spiral in Amborella and Austrobaileyales. It is unresolved whether phyllotaxis was ancestral...
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Context 1
... course of evolution of characters on two reference trees, chosen to represent the range of current phylogenetic results. One ("J/M") ( Fig. 4) corresponds to the chloroplast genome trees of Jansen & al. (2007) and Moore & al. (2007), where Chloranthaceae are sister to magnoliids and Ceratophyllum is sister to eudi- cots, while the other ("D&E") (Figs. 3, 4, 6) corresponds to the combined morphological and molecular analysis of Doyle & Endress (2000), where Chloranthaceae diverge at the base of the mesangiosperms, with the addition of Ceratophyllum as the sister group of Chloranthaceae and rearrangements within major clades based on more recent analyses. Additional results have further ...
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... optimization of perianth (and androecium) phyllotaxis shows that the ancestral character state in angio- sperms is equivocal, which is expressed by the hatched signa- ture at the base of the tree in Fig. 3 Doyle & Endress, 2011). In this analysis simple and complex whorled are lumped; the distinction between them is largely expressed by another character in the dataset, double positions. As with most characters considered in this article, the same ancestral state is found with both J/M and D&E refer- ence trees. Thus, we do not know ...
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... in which pollen tube growth and competition take place. Closing commonly begins at midlength of the style. At the lower end of the closure area, a short transverse slit may also form. Thus the entire line of closure has the shape of an inverted "T" or "Y". Fig. 4. A, most parsimonious course of evolution of carpel form on the D&E tree (see Fig. 3); B, evolution of carpel form on a tree with mesangiosperm rela- tionships based on nearly com- plete chloroplast genomes (J/M tree; Jansen & al., 2007;Moore & al., 2007). -Abbreviations as in Fig. 3 ...
Context 4
... entire line of closure has the shape of an inverted "T" or "Y". Fig. 4. A, most parsimonious course of evolution of carpel form on the D&E tree (see Fig. 3); B, evolution of carpel form on a tree with mesangiosperm rela- tionships based on nearly com- plete chloroplast genomes (J/M tree; Jansen & al., 2007;Moore & al., 2007). -Abbreviations as in Fig. 3 ...
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... before their key phylogenetic position was recognized (e.g., in Schisan- dra: Swamy, 1964;Cabomba Aubl.: Batygina & al., 1982;Galati, 1985;Nymphaea L.: Winter & Shamrov, 1991a;Van Miegroet & Dujardin, 1992;Orban & Bouharmont, 1998;Povilus & al., Fig. 6. Most parsimonious course of evolution of number of ovules per carpel on the D&E tree (see Fig. 3). In most groups with uniovulate carpels, the ovule direction is pendent; exceptions are indicated by grey arrows for ascendent direction, black arrow for horizontal direction. Abbreviations as in Fig. 3 ; Zini & al., 2015;Victoria: Winter & Shamrov, 1991a;Zini & al., 2015;Nuphar Sm.: Winter & Shamrov, 1991b;Barclaya: Winter, 1993). ...
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... 1992;Orban & Bouharmont, 1998;Povilus & al., Fig. 6. Most parsimonious course of evolution of number of ovules per carpel on the D&E tree (see Fig. 3). In most groups with uniovulate carpels, the ovule direction is pendent; exceptions are indicated by grey arrows for ascendent direction, black arrow for horizontal direction. Abbreviations as in Fig. 3 ; Zini & al., 2015;Victoria: Winter & Shamrov, 1991a;Zini & al., 2015;Nuphar Sm.: Winter & Shamrov, 1991b;Barclaya: Winter, 1993). However, the situation in Amborella is different. Tobe & al. (2000) reported that its embryo sac has three rounds of mitotic divisions, leading to a conventional Polygonumtype embryo sac, but later it was ...
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Citations
... Neue Untersuchungen (Scutt 2021) 146 gehen davon aus, dass ein "bract-derived perianth was likely present in the MRCA (= Most Recent Common Ancestor) of flowering plants (Endress and Doyle 2015) 147 may have shown a whorled arrangement (Sauquet et al. 2017) 148 . However perianth-like organs are not entirely specific to angiosperms as these were present in the flower-like reproductive axes of Bennettitales and also occur in the reproductive axes of all three extant genera of Gnetales: Ephedra, Gnetum and Welwitschia. ...
Homology, Type, basic organs, ontogeny, scale leaf, leaf, bud, flower, ontogeny, Angiosperms, methodology. Review.
Die Arbeit untersucht die erkenntnistheoretischen und methodischen Grundlagen der vergleichenden Morphologie. Die Begriffe Homologie und Analogie spielen eine zentrale Rolle. Teil der vergleichenden Morphologie ist die Organogenese. Die wichtigsten Konzepte sind das Homologie‐konzept, das Grundorgankonzept, das Typuskonzept. Die Arbeit enthält ein Review der verschiedenen Grundorgankonzepte und ein Review der Homologisierungen der Blüte mit dem Spross.
... Most angiosperm species are hermaphroditic, with individuals having both male and female sexual organs within the same flower. However, approximately 6% of angiosperm species are dioecious (Ricklefs, 1995;Renner, 2014), which are to have evolved from ancestral hermaphroditism (Endress and Doyle, 2015;Yakimowski and Barrett, 2016). Dioecious plants play an important role in protecting species diversity and maintaining ecosystem stability. ...
Dioecious plant species have a high genetic variation that is important for coping with or adapting to environmental stress through natural selection. Intensive studies have reported dimorphism morphism in morphology, physiology, as well as biotic and abiotic stress responses in dioecious plants. Here, we demonstrated the dimorphism of metabolic profile and the preference of some microorganisms in the roots and rhizosphere soils of male and female papaya. The metabolic composition of roots were significantly different between the males and females. Some sex hormones occurred in the differential metabolites in roots and rhizosphere soils. For example, testosterone was up-regulated in male papaya roots and rhizosphere soils, whereas norgestrel was up-regulated in the female papaya roots, indicating a possible balance in papaya roots to control the sexual differentiation. Plant hormones such as BRs, JAs, SA and GAs were also detected among the differential metabolites in the roots and rhizosphere soils of dioecious papaya. In addition, some metabolites that have medicinal values, such as ecliptasaponin A, crocin, berberine and sapindoside A were also expressed differentially between the two sexes. Numerous differential metabolites from the papaya roots were secreted in the soil, resulting in the differences in microbial community structure in the roots and rhizosphere soils. Some nitrogen-fixing bacteria such as Allorhizobium-Neorhizobium-Pararhizobium-Rhizobium, Brevundimonas and Microvirga were enriched in the male papaya roots or rhizosphere soils. While Candidatus Solibacter and Tumebacillus, which utilize organic matters, were enriched in the roots or rhizosphere soils of the female papaya. Some differences in the fungi abundance were also observed in both male and female papaya roots. These findings uncovered the effect of sex types on the metabolic and microbiota differences in roots and rhizosphere soils in papaya and will lead to investigations of underlining genomic and molecular mechanisms.
... All species of Ceratophyllum are specialized aquatics with underwater pollination [25,[39][40][41][42][43][44][45][46][47]. They lack any traces of roots. ...
... Homologies of structures bearing stamens and carpels in Ceratophyllum are problematic [25,39,40,[42][43][44][45][46][47][49][50][51]. Therefore, we use a neutral term 'reproductive unit' [50]. ...
... The reproductive units of Ceratophyllum were often interpreted as unisexual flowers with a perianth [41,52,53]. However, the idea of tepal homologies of involucral phyllomes is problematic because of the occasional occurrence of additional reproductive unit(s) inside the involucre [40,43,47]. According to Endress [51] and Endress and Doyle [47], each stamen of Ceratophyllum should be viewed as an individual male flower possessing no subtending bract. ...
Ceratophyllum is an ancient and phylogenetically isolated angiosperm lineage. Comparisons between Ceratophyllum and other angiosperms are hampered by uncertainty in inferring organ homologies in this genus of specialized aquatics. Interpretation of shoot morphology is especially problematic in Ceratophyllum. Each node has several leaf-like appendages interpreted as verticillate leaves, modified parts of one and the same leaf or parts of two leaves under decussate phyllotaxis. Vegetative branches are axillary, but reproductive units (interpreted as flowers or inflorescences) are commonly viewed as developing from collateral accessory buds. We studied shoot development in Ceratophyllum submersum, C. tanaiticum, and C. demersum using scanning electron microscopy to clarify shoot morphology and branching patterns. Our data support the idea that the phyllotaxis is essentially decussate with appendages of stipular origin resembling leaf blades. We conclude that a leaf axil of Ceratophyllum possesses a complex of two serial buds, the lower one producing a vege-tative branch and the upper one developing a reproductive unit. The reproductive unit is congenitally displaced to the subsequent node, a phenomenon known as concaulescence. Either member of the serial bud complex may be absent. There is a theory based on a synthesis of molecular and morphological data that Chloranthaceae are the closest extant relatives of Ceratophyllum. Serial buds and concaulescence are known in Hedyosmum (Chloranthaceae). Our new interpretation facilitates morphological comparisons between Hedyosmum and Ceratophyllum.
... Throughout the course of floral evolution in angiosperms, it has been considered that stability in floral merisms is correlated with occurrence of whorled phyllotaxis (Endress 2010a). Along with whorled phyllotaxis and radial symmetry, a small and stable merism is considered a prerequisite for the evolution of complex synorganizational patterns in flowers (Endress 2010a(Endress , 2016Endress and Doyle 2015). It is important to highlight the possible link between merism lability and meristem size and/or space. ...
Floral features contribute with remarkable additions to morphological studies and are widely used to address questions about the evolution and diversification of several groups of plants. Even though Simaroubaceae are a small monophyletic family, the few detailed structural analyses of reproductive organs and the floral diversity and variations already described in their members stimulate novel structural studies. In this study, we investigate the evolution of reproductive features of Simaroubaceae by means of a combination of original data and a review of the literature, aiming to elucidate which floral characters are most informative for a better understanding of the evolutionary history of the group. We analyzed 21 out of the 23 genera of Simaroubaceae, plus six from Rutaceae and seven from Meliaceae as outgroups. We used a Bayesian method and the Parsimony optimality criterion to reconstruct ancestral reproductive character states using a re-analyzed phylogenetic tree of Sapindales. Here, we combined available molecular sequences to have the largest sample of Simaroubaceae genera. We found that the ancestral flowers of Simaroubaceae were probably polygamous or dioecious plants, with free carpels united only distally, with divergent, elongated stigmas, and with drupaceous, laterally flattened to lenticular fruits. The latter feature plus apocarpous carpels are putative synapomorphies of the family retrieved in this study. Imbricate petals and a diplostemonous androecium were recovered as conditions found in the ancestor of Simaroubaceae but also shared with the ancestors of Meliaceae and Rutaceae. Our findings were mostly in accordance with previous evolutionary studies on genera of Simaroubaceae and with other families of Sapindales. © 2021, The Author(s), under exclusive licence to Botanical Society of Sao Paulo.
... The latest and most comprehensive study [1] concluded that the ancestral angiosperm flower had multiple free carpels which were simple, with distal openings closed by secretion. This result agrees with previous studies by Endress and Doyle [2,3] who have built a wealth of knowledge on the morphology of flowering plants and the reproductive Fig. 1 Schematic representation of the carpel and phylogenetic relationships of seed plants. A The carpel consists of an ovary which produces the ovules, the stigma which receives the pollen and the style which guides the pollen tube to the ovules. ...
The carpel is a fascinating structure that plays a critical role in flowering plant reproduction and contributed greatly to the evolutionary success and diversification of flowering plants. The remarkable feature of the carpel is that it is a closed structure that envelopes the ovules and after fertilization develops into the fruit which protects, helps disperse, and supports seed development into a new plant. Nearly all plant-based foods are either derived from a flowering plant or are a direct product of the carpel. Given its importance it’s no surprise that plant and evolutionary biologists have been trying to explain the origin of the carpel for a long time. Before carpel evolution seeds were produced on open leaf-like structures that are exposed to the environment. When the carpel evolved in the stem lineage of flowering plants, seeds became protected within its closed structure. The evolutionary transition from that open precursor to the closed carpel remains one of the greatest mysteries of plant evolution. In recent years, we have begun to complete a picture of what the first carpels might have looked like. On the other hand, there are still many gaps in our understanding of what the precursor of the carpel looked like and what changes to its developmental mechanisms allowed for this evolutionary transition. This review aims to present an overview of existing theories of carpel evolution with a particular emphasis on those that account for the structures that preceded the carpel and/or present testable developmental hypotheses. In the second part insights from the development and evolution of diverse plant organs are gathered to build a developmental hypothesis for the evolutionary transition from a hypothesized laminar open structure to the closed structure of the carpel.
... The conflicting evolutionary relationships among magnoliids, monocots, and eudicots also reflect the complexity of their morphologies, especially floral organs (Endress and Doyle, 2015). The flowers of C. camphora show a gradual morphological transition from being sepal-like to petal-like. ...
Magnoliids represent the third largest branch of the most diverse and species-rich angiosperms, and their phylogenetic position relative to monocots and eudicots remains uncertain. Here, we describe the assembly of a high-quality chromosome-level genome sequence for Cinnamomum camphora (camphor), an evergreen tree belonging to Lauraceae. Genome-based phylogenetic analysis indicated possible incomplete lineage sorting (ILS) during the rapid diversification of the early diverging branches of angiosperms, which may have led to the unstable phylogenetic position of magnoliids. Whole-genome duplication (WGD) analysis showed that the lineage leading to C. camphora experienced two WGD events: the earlier event was shared by the common ancestor of Chimonanthus salicifolius and Lauraceae, and the more recent event was shared by all lineages of Lauraceae. The ABCDE homolog genes of C. camphora showed broad expression in floral organs, revealing the molecular mechanism preventing tepals from differentiating into petals and sepals. The key genes for the synthesis of monoterpenes have been identified, and the molecular regulatory mechanism underlying the rich essential oils in the branches and leaves has been revealed. Overall, our study provides a better understanding of the evolution and diversification of magnoliids and the evolution of floral organs.
... Floral variation is a positive response of plants to the selection pressure [38][39][40][41]. Studies on floral variation not only contributed to our understanding of species evolution [42][43][44][45], but also revealed the genetic rules and variation degrees of populations / groups [46][47][48][49][50], which in turn provided a theoretical basis for the protection of species. In this study, through the cluster analysis of the 142 Malus taxa based on their floral organs phenotypic traits, we found that the distribution of Malus species belonged to the same section (series) was relatively concentrated, with ancestor-inclined distribution probability reaching up to 87% in two groups (A, B) and 61% in five sub-groups (A 1 , A 2 , B 1 , B 2 , B 3 ). ...
... In fact, the variation or change in different floral organs usually occurs at different taxonomic levels (family, genus, species, ranks below species) [52]. Size, color, smell, and the taste of floral organs are often quite different in species or lower levels [45,[53][54][55]. Jin [56] concluded that in the taxonomy of subgenus Tsutsusz (Rhododendron), the tree habit, shape and the size of corollas, could be used to distinguish grades above the species level. ...
Background
In angiosperms, phenotypic variation of floral organs is often considered as the traditional basis for the evolutionary relationship of different taxonomic groups above the species level. However, little is known about that at or below the species level. Here, we experimentally tested the phenotypic variation of Malus floral organs using combined methods of intraspecific uniformity test, interspecific distinctness analysis, principal component analysis, Pearson correlation analysis, and Q-type cluster analysis. The ancestor-inclined distribution characteristic analysis of Malus species and cultivars floral attributes was also carried out, so as to explore its taxonomic significance.
Results
15/44 phenotypic traits (e.g., flower shape, flower type, flower diameter, ...) were highly consistent, distinguishable, and independent and could be used as the basis for Malus germplasm taxonomy. The studied 142 taxa were divided into two groups (A, B) and five sub-groups (A 1 , A 2 , B 1 , B 2 , B 3 ), with significantly variable floral phenotypic attributes between groups and within sub-groups. Malus natural species were relatively clustered in the same section (series) while homologous cultivars showed evidence of ancestor-inclined distribution characteristics. However, no significant correlation between the evolutionary order of sections (Sect. Docyniopsis → Sect. Chloromeles → Sect. Sorbomalus → Sect. Eumalus ) and group/sub-groups (B 3 → B 2 → B 1 → A).
Conclusions
Phenotypic variation of floral organs could better explore the genetic relationship between Malus taxa. The findings improved our cognition of floral phenotypic variation taxonomic significance under the species level.
... Apparently, the synascidiate zone is short in this species, and it is completely sterile, i.e. the placentas are restricted to the symplicate zone and initially entirely parietal (without an axile part). Completely plicate carpels are quite rare among monocots and angiosperms in general (Igersheim, Buzgo & Endress, 2001;Remizowa, Sokoloff & Rudall, 2010;Endress & Doyle, 2015). In Dioscoreales, completely plicate carpels have been reported for Tacca J.R.Forst. ...
Thismia is characterized by an exceptionally complicated floral morphology that is currently not understood properly. In the taxonomic literature, descriptive rather than morphological terms are often applied to parts of the flower in Thismia, relating to the general appearance of the floral organs instead of their precise homologies. Precise understanding of the floral structure is complicated by the rarity of Thismia spp. and the paucity of appropriate material. Here we provide a comprehensive study of reproductive organs of three Thismia spp. (T. annamensis, T. javanica and T. mucronata) including the first investigation of inflorescence architecture and early floral development in Thismiaceae. We found a hitherto unknown diversity of the reproductive shoots in the genus, manifested in the number of floral prophylls (two or three, in contrast to a single prophyll in the vast majority of monocots) and in the branching plane resulting in two distinct inflorescence types, a drepanium and a bostryx. We report the non-acropetal sequence of initiation of floral whorls (with stamens being the last elements to initiate), never previously described in monocots, and the gynoecium composed of completely plicate carpels, also a rare feature for monocots. Floral vasculature is relatively uniform in Thismia, but significant interspecific differences are found in tepal innervation, including the number of tepal traces; some of these differences are not immediately related to the external tepal morphology. We argue that the annulus, which acts as a roof of the hypanthium, possesses an androecium nature and represents congenitally fused bases of stamen filaments. We describe the stamens as laminar structures, which are also shortly tubular in the distal part of the supraconnective with the adaxial tubular side forming a skirt-like appendage. Finally, the placentas, which are column-like when mature, are initially parietal, becoming secondarily similar to free-central placentas through schizogenous separation from the ovary wall.
... A synstigma is a rarely reported structure of angiosperms formed by the clustering of two or more stigmas originating from a single flower or from different flowers. Records of synstigmas have been reported in: early-diverging angiosperms (Amborellaceae, Austrobaileyaceae, Illiciaceae, Nymphaeaceae, Schisandraceae) and magnoliids (Annonaceae, Calycanthaceae, Monimiaceae, Siparunaceae), in which the gynoecium is apocarpic, but contacted by an extragynoecial compitum (Endress, 2011;Endress & Doyle, 2015); Rosa setigera Michx. (Rosaceae), in which the apocarpic gynoecium is contacted by an intrastylar compitum (Kemp, Kevan & Posluszny, 1993); for two species of Procris Comm. ...
... Thus, the synstigma of Ficus is functionally analogous to a special type of compitum, the extragynoecial found in some groups, formed in the external surfaces of the gynoecia (Endress, 1979(Endress, , 1982Endress & Lorence, 1983;Endress & Igersheim, 2000;Vieira & Shepherd, 2002;Lyew et al., 2007;Du & Wang, 2012;Lau et al., 2017). A common type of extragynoecial compitum involves abundant mucilage secreted by the stigmatic papillae, as commonly found in some groups of earlydiverging angiosperms (Endress, 1979(Endress, , 2011Endress & Lorence, 1983;Endress & Igersheim, 2000;Wang, Armbruster & Huang, 2012;Endress & Doyle, 2015;Lau et al., 2017). The mucilage serves as a germination medium and aids in the selection and distribution of pollen tubes between the carpels. ...
The synstigma is a structure formed by clusters of two to several stigmas, whether in the same or between different flowers. Although rare in angiosperms, synstigmas are found in c. 500 out of the c. 750 Ficus spp. (Moraceae). This floral structure is associated with fig-fig wasp pollinating mutualism. The synstigma structure and pollen tube pathways were studied in six Ficus spp. from Ficus section Americanae to test the hypothesis that the synstigma allows pollen grains deposited on a stigma to emit pollen tubes that can grow laterally and fertilize surrounding flowers. Syconia containing recently pollinated stigmas were collected and dissected, and the stigmas were processed for analyses with light and scanning and transmission electron microscopy. The arrangement of the synstigmas across species can be spaced or congested, with the number of stigmas per synstigma ranging from two to 20. Contact between the stigmas in a synstigma occurs by the intertwining of the stigmatic branches and papillae; their union is firm or loose. The pollen tube grows through live cells of the transmitting tissue until reaching the ovule micropyle. Curved pollen tubes growing from one stigma to another were observed in five out of the six species studied. The curvilinear morphology of pollen tubes probably results from competition by pollen between the stigmas composing a synstigma via chemotropic signals. The synstigma appears to be a key adaptation that ensures seed production by flowers not exploited by the fig wasps in actively pollinated Ficus spp. ADDITIONAL KEYWORDS: active pollination-Ficus-Moraceae-pollen grain-pollen-pistil interaction-pollen tube-ultrastructure.
... However, we cannot completely rule out this possibility, as frequent sex chromosome turnover is possible in some 30 lineages (37,38). The structure of the A. trichopoda flower, however, does not fit well with ancient dioecy, as non-functional male organs are present in female flowers, while a pronounced central dome in male flowers may be a remnant of the gynoecium (39). Overall, our results tend to support the idea that the most recent ancestor of all extant angiosperms was indeed bisexual. ...
Sex determination is poorly understood in plants. Amborella trichopoda is a well-known plant model for evo-devo studies, which is also dioecious (has male and female individuals), with an unknown sex determination mechanism. A. trichopoda is a “sex switcher”, which points to possible environmental factors that act on sex, but populations grown from seed under greenhouse conditions exhibit a 50:50 sex ratio, which indicates the operation of genetic factors. Here, we use a new method ( SDpop ) to identify sex-linked genes from genotyping data of male and female individuals sampled in the field, and find that A. trichopoda has a ZW sex-chromosome system. The sex-linked genes map to a 4 Mb sex-determining region on chromosome 9. The low extent of ZW divergence suggests these sex chromosomes are of recent origin, which is consistent with dioecy being derived character in the A. trichopoda lineage. Our work has uncovered clearly formed sex chromosomes in a species in which both genetic and environmental factors can influence sex.
One Sentence Summary
Amborella trichopoda , a dioecious species in which both genetics and the environment influence sex, possesses a pair of quite recently evolved ZW chromosomes.
