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Seed morphology in Brassicales: configurations of embryo and testa. Cleomaceae (A cl ) and Brassicaceae (A-G), with testa indicated by short parallel lines.-A cl. Cleomaceae, all species have uniformly incumbent cotyledons, e.g., Tarenaya hassleriana (Chodat) Iltis (¼ Cleome hassleriana Chodat) (cf. Fig. 2I-W, Cleomaceae; Fig. 2P, T. hassleriana; Iltis, Cochrane & Tucker, unpub.). A-G. Brassicaceae, showing the great diversity of embryo and testa configurations in this family, cf. Fig. 2X-Z.-A. Sisymbrium strictissimum L., with incumbent cotyledons.-B. Cheiranthus cheiri L., accumbent cotyledons.-C. Brassica nigra (L.) W. D. J. Koch, conduplicate cotyledons.-D. Chamira circaeoides (L. f.) Zahlbr., doubly conduplicate cotyledons.-E. Calepina irregularis (Asso) Thell., cross-folded cotyledons.-F. Bunias erucago L., spirolobous cotyledons.-G. Heliophila pusilla L. f., double cross-folded cotyledons. A cl , from Iltis original drawing; A-G, adapted from diagrams in Melchior (1964: 184).
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Molecular data support the recognition of three monophyletic families, Capparaceae s. str., Cleomaceae, and Brassicaccae, instead of an all-encompassing Brassicaceae or a paraphyletic Capparaceae s.l. This view is reinforced with many figures showing two basic and ubiquitous differences in cleomoid seed structure. First, the more or less strongly i...
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Citations
... The gynophore of Arachis hypogaea (peanut) in Fabaceae is an exception because it has been more thoroughly researched due to its essential role in underground fruit development (Moctezuma, 2003;Xia et al., 2013;Zhao et al., 2015); however, this gynophore is unique in that its formation is initiated after fertilization. Though androgynophores and gynophores that develop during flowering are often inconspicuous, prominent stalk-like structures are characteristic of Cleomaceae (Iltis et al., 2011) and are present in other distantly related species, including the crop plant Passiflora edulis (passion fruit) in Passifloraceae. ...
Premise:
Flowering plants have evolved a vast array of floral features involved in plant-pollinator interactions. A feature that seemingly increases the chance of pollen transfer is the androgynophore; a stalk-like structure that raises the reproductive organs of the flower. However, little is known about the developmental and genetic basis of this structure despite its presence in multiple, distantly related taxa. Here, we address this gap by investigating Gynandropsis gynandra (Cleomaceae), a species with prominent androgynophore.
Methods:
We combined morphological and anatomical analyses with a comparative transcriptomic study to provide a detailed description of the androgynophore throughout development, examine global gene expression patterns, and identify candidate genes putatively involved in androgynophore elongation.
Key results:
The radially symmetric androgynophore of G. gynandra rapidly lengthens primarily via cell elongation. Despite its structural uniformity, androgynophore development is characterized by complex gene expression patterns including differential expression of floral organ identity genes and genes associated with organ development and growth in Arabidopsis thaliana.
Conclusions:
Our morphological characterizations and high-quality transcriptome for G. gynandra suggest that the androgynophore is a novel structure formed via elaboration of both the receptacle and base of reproductive organs as it is structurally like an elongated internode but expresses the genetic repertoire typically associated with the reproductive organs. The drastic increase in cell length and uniform structure elevates the androgynophore as a potentially powerful model for cell elongation. This article is protected by copyright. All rights reserved.
... The family is distributed worldwide, comprising 26 genera and 270 species (Bayat et al., 2018;Stevens, 2001;Soares-Neto et al., 2020, 2022. Cleomaceae can be easily distinguished from other plant families based on specific morphological traits, such as compound leaves (3-12 leaflets), zygomorphic flowers, six stamens, and dry fruits with longitudinal dehiscence (Stevens, 2001;Iltis et al., 2011). Members of this family are used as model plants in studies of floral evolution, C 4 photosynthesis, and comparative genomics and transcriptomics (Brown et al., 2005;Feodorova et al., 2010;Nozzolillo et al., 2010). ...
... The species studied here reflect the remarkable diversity of the Cleomaceae family in terms of morphology (Patchell et al., 2011), photosynthetic mechanisms (Feodorova et al., 2010;Marshall et al., 2007;Voznesenskaya et al., 2007), genomics (Inda et al., 2008), pollination (Cane, 2008;Machado et al., 2006;Zohoungbogbo et al., 2018), and geographic distribution (Bayat et al., 2018). Accordingly, this family has been proposed as a model thanks to its several peculiarities and its close phylogenetic relationship with Arabidopsis thaliana of the Brassicaceae family (Iltis et al., 2011), from which Cleomaceae has diverged relatively recently (~35 mya.) (Schranz and Mitchell-olds, 2006). ...
... Gynandropsis gynandra (2n = 34, shares the common name "spider plant" with a number of unrelated species) belongs to the Cleomaceae, the sister family of the Brassicaceae (Hugh et al., 2011), and is grown as a leafy vegetable but also as a medicinal plant (Sogbohossou et al., 2018). Gynandropsis gynandra is an essentially cosmopolitan species found across Africa, Asia, the Middle East, and Australasia and has been introduced to the Caribbean, Southern and Northern America, and Central and Northern Europe (Chweya and Mnzava, 1997). ...
Gynandropsis gynandra (Cleomaceae) is a cosmopolitan leafy vegetable and medicinal plant, which has also been used as a model to study C4 photosynthesis due to its evolutionary proximity to C3 Arabidopsis (Arabidopsis thaliana). Here, we present the genome sequence of G. gynandra, anchored onto 17 main pseudo-molecules with a total length of 740 Mb, an N50 of 42 Mb and 30,933 well-supported gene models. The G. gynandra genome and previously released genomes of C3 relatives in the Cleomaceae and Brassicaceae make an excellent model for studying the role of genome evolution in the transition from C3 to C4 photosynthesis. Our analyses revealed that G. gynandra and its C3 relative Tarenaya hassleriana shared a whole-genome duplication event (Gg-α), then an addition of a third genome (Th-α, + 1x) took place in T. hassleriana but not in G. gynandra. Analysis of syntenic copy number of C4 photosynthesis-related gene families indicates that G. gynandra generally retained more duplicated copies of these genes than C3T. hassleriana, and also that the G. gynandra C4 genes might have been under positive selection pressure. Both whole-genome and single-gene duplication were found to contribute to the expansion of the aforementioned gene families in G. gynandra. Collectively, this study enhances our understanding of the polyploidy history, gene duplication and retention, as well as their impact on the evolution of C4 photosynthesis in Cleomaceae.
... DeWolf is a perennial woody herb or a subshrub belonging to the Capparaceae or caper family. The Capparaceae family is a tropical and subtropical taxon of trees, shrubs, herbs and lianas consisting of approximately 417 species (Kers and Capparaceae, 2003;Iltis et al., 2011). The genus Maerua Forssk. ...
A Maerua pseudopetalosa (Gilg & Gilg-Ben.) DeWolf is a perennial herb or sub-shrub collected from the wild as a food plant or traditional medicine. Maerua pseudopetalosa occurs naturally from Senegal, east to Eritrea and Ethiopia, through South Sudan, Sudan, East Africa, south to the Democratic Republic of Congo (DRC). This study is aimed at evaluating the chemical properties, biological activities and ethnomedicinal uses of M. pseudopetalosa. Results of the current study are based on data derived from several online databases such as Scopus, Google Scholar, PubMed and Science Direct, and pre-electronic sources such as scienti ic publications, books, dissertations, book chapters and journal articles. This study revealed that the fruit, leaf and root infusion and/or decoction of M. pseudopetalosa are mainly used as a stimulant and to restore physical strength and vitality, and traditional medicine for chest pains, breast cancer and cough. Phytochemical compounds identi ied from aerial parts, fruits, roots, seeds and tubers of M. pseudopetalosa include quater-nary ammonium compounds, alkaloids, amino acids, betaines, cardiac glyco-sides, fatty acids, essential oils, lavonoids, phenolics, sterols and tannins. Preliminary ethnopharmacological research revealed that aerial parts, roots and tubers of M. pseudopetalosa have antibacterial, antifungal, antihyperglycemic, antioxidant, α-glucosidase inhibitory, pancreatic lipase inhibitory and cyto-toxicity activities. Maerua pseudopetalosa should be subjected to detailed phy-tochemical, pharmacological and toxicological evaluations aimed at correlating its medicinal uses with its phytochemistry and pharmacological properties .
... Maerua triphylla A. Rich. is a shrub or small tree belonging to the Capparaceae or caper family. The Capparaceae family is a tropical and subtropical taxon of trees, shrubs, herbs and lianas consisting of approximately 417 species (Kers et al., 2003;Iltis et al., 2011) The genus Maerua Forssk has been recorded in tropical and southern Africa, and tropical Asia with about 60 species recorded from the African continent (Elfers et al., 1964). The genus name "Maerua" means "drooping" about the drooping foliage (Venter and Venter, 2015). ...
... The genus Cleome and other genera of the family have been traditionally considered as the subfamily Cleomoïdeae of the family Capparaceae (Inda et al., 2008). In recent phylogenetic studies, Cleomoïdeae were found to be more closely related to the Brassicaceae than Capparoideae (Hall, 2008;Hall et al., 2002;Iltis et al., 2011). Thus, three monophyletic families were delineated: Capparaceae, Cleomaceae, and Brassicaceae ( Figure 1). ...
... The morphological Capparaceae Arivela (10) Cleome (200) Cleomella (10) Cleoserrata (5) Dactylaena (6) Dipterygium (1) Gynandropsis (1) Hemiscola (6) Oxystylis (1) Peritoma (6) Physostemon (17) Podandrogyne (36) characters specific to the Cleomaceae include their mostly herbaceous habit, compound leaves and dry and dehiscent capsules with a replum. Seeds are uniformly strongly curved, reniform or horseshoe-shaped to conduplicate with a deep invagination of the testa, an incurved embryo with always strictly incumbent cotyledons (Iltis et al., 2011). Gynandropsis gynandra was also known as Cleome gynandra L. (1753), Cleome pentaphylla L. (1763), and G. pentaphylla (L.) DC. (1824). ...
This article reveals progress and gaps in spider plant (Gynandropsis gynandra Syn. Cleome gynandra) research to date. It synthesises the current knowledge in systematics and botany, geographical distribution, nutrient content, and health values as well as the agronomic practices related to the species. This article further presents the germplasm availability, status, and places of conservation around the world to exploit the extant genetic diversity for genomics‐assisted breeding and seed systems development. It describes how various parts of the plant have been used and the potential for economical revamping and further industrial valorisation. Nutrient content and phytochemical diversity across cultivation systems and geographical ranges and known health attributes were analysed to identify gaps and bottlenecks for fostering further investigations for industrial food and drug production.
... Also, like Brassicaceae, fruits of C. violacea have a prominent persistent placenta called the replum. The primary difference between Cleomaceae capsule fruits and Brassicaceae siliques is that Cleomaceae fruits lack the false septum separating the two locules ( Fig. 2B) (Iltis et al., 2011). Like with PDS, initial experiments with pTRV2-CvFUL-CvPDS were completed to determine effectiveness in C. violacea. ...
Premise:
Cleomaceae is emerging as a promising family to investigate a wide range of phenomena, such as C4 photosynthesis and floral diversity. However, functional techniques are lacking for elucidating this diversity. Herein, we establish virus-induced gene silencing (VIGS) as a method of generating functional data for Cleome violacea, bolstering Cleomaceae as a model system.
Methods:
We leveraged the sister relationship of Cleomaceae and Brassicaceae by using constructs readily available for Arabidopsis thaliana to provide initial information about the feasibility of VIGS in C. violacea. We then developed endogenous constructs to optimize VIGS efficiency and viability for fruit development.
Results:
PHYTOENE DESATURASE was successfully downregulated in C. violacea using both heterologous and endogenous constructs. The endogenous construct had the highest degree of downregulation, with many plants displaying strong photobleaching. FRUITFULL-treated plants were also successfully downregulated, with a high rate of survival but less effective silencing; only a small percentage of survivors showed a strong phenotype.
Discussion:
Our optimized VIGS protocol in C. violacea enables functional gene analyses at different developmental stages. Additionally, C. violacea is amenable to heterologous knockdown, which suggests that a first pass using non-endogenous constructs is a possible route to test additional species of Cleomaceae.
... Згідно з даними Angiosperm Phylogeny Group (APG IV, 2016), Cleomaceae є сестринською родиною Brassicaceae. У видів клеомових численні й дуже різноманітні за структурою залозисті волоски й рідкісні або зовсім відсутні прості трихоми (Pax, 1891;Iltis et al., 2011;Ilyinska, 2015). Отже, розвиток залозистих структур також вказує на існування близьких споріднених зв'зків між родинами Brassicaceae та Cleomaceae і є одним із свідчень паралельного розвитку цих двох родин (Hall et al., 2002;Iltis et al., 2011;APG-IV, 2016). ...
... У видів клеомових численні й дуже різноманітні за структурою залозисті волоски й рідкісні або зовсім відсутні прості трихоми (Pax, 1891;Iltis et al., 2011;Ilyinska, 2015). Отже, розвиток залозистих структур також вказує на існування близьких споріднених зв'зків між родинами Brassicaceae та Cleomaceae і є одним із свідчень паралельного розвитку цих двох родин (Hall et al., 2002;Iltis et al., 2011;APG-IV, 2016). У таксонів флори України, що належать до LIII, просте опушення складається із нерозгалужених (Clausia та Chorispora) або із простих та фуркатних (триби Hesperideae, Euclidieae та Buniadeae) чи із зірчастих і дендроїдних (Anchonieae) трихом. ...
Species of the Brassicaceae family (crucifers), which are of great scientific and practical importance, are inherent by a wide variety of trichome structures. Trichomes – specialized cells of the epidermis, modulate the response of plants to biotic and abiotic stresses. We investigated the structural diversity of trichomes and the features of pubescence of all (252 species, 73 genera) Brassicaceae species of the flora of Ukraine. Trichomes were studied on living plants and based on materials from seven herbaria, as well as on the basis of analysis of literature data. Stereoscopic and electronic scanning (JSM-6060 LA) microscopes were used in this investigation. To describe the structural diversity of trichomes, an original four-level hierarchical classification was developed, taking into account a complex of features, including functional specificity (covering or glandular), the nature and degree of hair branching, as well as the features of the orientation of their rays. The pubescence of plants was divided into isomorphic (consists of one type of trichomes) and heteromorphic (formed by two or more types of hairs with or without glandular structures). Hairs are completely absent in 23 species of 15 genera of the flora of Ukraine. Isomorphic pubescence is characteristic of the most (170 species) cruciferous plants and is represented by simple (120 species), Malpighian (23 species), stellate (23 species), dendroid (3) and fork-shaped (1 species) trichomes. The composition of heteromorphic pubescence is dominated by a combination of simple and fork-shaped hairs: it is observed in 44 species. There are glandular structures in the pubescence of 14 species. The structural diversity of Brassicaceae trichomes correlates, in general, with the temperate continental climate of Ukraine. Species without hairs or pubescent with simple trichomes, including those in combination with fork-shaped hairs, are observed in biotopes with sufficient moisture, optimal insolation, and satisfactory soil quality. Species with dense branched trichomes inhabit arid biotopes with intense
insolation, insufficient moisture, and poor substrate. In the flora of Ukraine, the evolutionary-phylogenetic line LII included non-pubescent species or those with simple hairs. The cruciferous line LI is characterized by isomorphic and heteromorphic pubescence. In species LIV, heteromorphic pubescence predominates. Glandular structures are observed in the heteromorphic pubescence of taxa of line LIII.
... Morphotype 5 (Fig. 33I-K) is interpreted as the cast of a strongly curved, reniform, or horseshoe-shaped seed with an incurved embryo and is identified as aff. Cleomaceae, whose seeds have strongly incurved embryos and a deep invagination of the testa (Iltis et al. 2011). Morphotype 7 (Fig. 33M1-O) is identified as aff. ...
The Pliocene–Pleistocene transition in the Neotropics is poorly understood despite the major climatic changes that occurred at the onset of the Quaternary. The San Gregorio Formation, the younger unit of the Urumaco Sequence, preserves a fauna that documents this critical transition. We report stingrays, freshwater bony fishes, amphibians, crocodiles, lizards, snakes, aquatic and terrestrial turtles, and mammals. A total of 49 taxa are reported from the Vergel Member (late Pliocene) and nine taxa from the Cocuiza Member (Early Pleistocene), with 28 and 18 taxa reported for the first time in the Urumaco sequence and Venezuela, respectively. Our findings include the first fossil record of the freshwater fishes Megaleporinus, Schizodon, Amblydoras, Scorpiodoras, and the pipesnake Anilius scytale, all from Pliocene strata. The late Pliocene and Early Pleistocene ages proposed here for the Vergel and Cocuiza members, respectively, are supported by their stratigraphic position, palynology, nannoplankton, and 86Sr/88Sr dating. Mammals from the Vergel Member are associated with the first major pulse of the Great American Biotic Interchange. In contrast to the dry conditions prevailing today, the San Gregorio Formation documents mixed open grassland/forest areas surrounding permanent freshwater systems, following the isolation of the northern South American basin from western Amazonia. These findings support the hypothesis that range contraction of many taxa to their current distribution in northern South America occurred rapidly during at least the last 1.5 million years.
... Plumbaginaceae and Zygophyllaceae, which represent quite a few halophytes, for example usually have well-developed straight or curved embryos (Corner, 1976). In Brassicaceae, however, some species are known to germinate very fast as well (Parsons, 2012), while spirally coiled embryos are known to occur in some ephemeral species (Iltis et al., 2011). Nonetheless, more studies are required to assess seed morphology and ecology relationships in other plant families in general and in halophytes specifically. ...
Background and aims:
Internal seed morphological traits such as embryo characteristics and nutritive tissue can vary considerably within a plant lineage. These traits play a prominent role in germination processes and the success of seedling establishment, and are therefore under high selective pressure, especially in environments hostile to seedlings such as arid, saline or highly dynamic habitats. We investigated the relationships of seed internal morphology and germination characteristics of 84 species of Amaranthaceae s.l., a family with numerous lineages that have adapted to stressful growing conditions.
Methods:
We used seed cross sections to assess embryo type, embryo to seed surface ratio and radicle to cotyledon length ratio. Furthermore, seed mass, mean time to germination, habitat preferences and further plant traits such as C3 or C4 photosynthesis and life form were compiled for each species. Data were analyzed using phylogenetic comparative methods.
Key results:
We found embryo type (λ = 1), log seed mass (λ = 0.86) and embryo to seed size ratio (λ = 0.78) to be evolutionarily stable with an annular embryo as ancestral in the family. Linked to shifts to the three derived embryos types (spiral, horseshoe-shaped and curved) is an increase in root to cotyledon length ratio and a reduction of nutritive tissue. We observed stabilizing selection towards seeds with relatively larger embryos with longer radicles and less nutritive tissue that are able to germinate faster, especially in lineages with C4 photosynthesis and/or salt tolerance.
Conclusions:
We conclude that the evolutionary shift of nutrient storage from perisperm to embryo provides an ecological advantage in extreme environments, because it enables faster germination and seedling establishment. Furthermore, the evolutionary shift towards higher root to cotyledon length ratio especially in small-seeded Amaranthaceae growing in saline habitats can provide an ecological advantage for fast seedling establishment.
