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Distribution of the seven Psidium species in Brazilian biomes based on data from the literature, herbaria, and the present study. Distribution of the diploid species (2n = 2x = 22 chromosomes): P. cauliflorum (blue circle), P. guajava (black circle), and P. oblongatum (yellow circle). Note that P. cauliflorum and P. oblongatum (yellow circle) only occur in the Atlantic Forest. Tetraploid species (2n = 4x = 44 chromosomes): distribution of P. guineense (black square) was not registered in Pantanal and the Pampas. Hexaploid species (2n = 6x = 66 chromosomes): P. cattleyanum (green triangle) and P. myrtoides (red triangle) occur in the Atlantic Forest, Cerrado, Caatinga, and the Pampas. Octoploid species (2n = 8x = 88 chromosomes): P. longipetiolatum (blue hexagon) is found distributed in the Atlantic Forest from Minas Gerais to Rio Grande do Sul. The coordinates used to define the species’ geographical distribution were obtained with the application Google Earth, using the locations indicated on the labels of the herbarium specimens. The map of geographical distribution was made using the program DIVA GIS 5.4

Distribution of the seven Psidium species in Brazilian biomes based on data from the literature, herbaria, and the present study. Distribution of the diploid species (2n = 2x = 22 chromosomes): P. cauliflorum (blue circle), P. guajava (black circle), and P. oblongatum (yellow circle). Note that P. cauliflorum and P. oblongatum (yellow circle) only occur in the Atlantic Forest. Tetraploid species (2n = 4x = 44 chromosomes): distribution of P. guineense (black square) was not registered in Pantanal and the Pampas. Hexaploid species (2n = 6x = 66 chromosomes): P. cattleyanum (green triangle) and P. myrtoides (red triangle) occur in the Atlantic Forest, Cerrado, Caatinga, and the Pampas. Octoploid species (2n = 8x = 88 chromosomes): P. longipetiolatum (blue hexagon) is found distributed in the Atlantic Forest from Minas Gerais to Rio Grande do Sul. The coordinates used to define the species’ geographical distribution were obtained with the application Google Earth, using the locations indicated on the labels of the herbarium specimens. The map of geographical distribution was made using the program DIVA GIS 5.4

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Key message Polyploidy (diploid to octoploid) was evidenced from seven Psidium species, besides the outcomes of the whole-genome duplication about the nuclear DNA content, DNA sequence, and distribution. Abstract The previous studies have reported the occurrence of polyploid species in Psidium, all deriving from the basic chromosome number x = 11,...

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... Among N. sibirica plants, the highest variation of genome size (1.02-fold) was found in populations of Altai Krai (Uglovskoe), Altai Republic (Kosh-Agach), Tuva (Turan), and Kazakhstan (Koktal), located at a substantial distance from each other. Correlations between ecological differentiation and genome size have been reported at both interspecific and intraspecific levels (Knight et al. 2005;Knight and Ackerly 2002), for example, in plant species of genera Larrea (Poggio et al. 1989), Cardiospermum (Urdampilleta et al. 2012), Berberis (Bottini et al. 2000), Coffea (Razafinarivo et al. 2012), and Psidium (Tuler et al. 2019). For instance, it was shown that the species of Larrea, Bulnesia, and Pintoa that inhabit the most arid environments are the ones possessing the highest DNA content (Poggio et al. 1989). ...
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For the first time, nuclear genome size and ploidy of five Nitraria species from 49 populations were examined by flow cytometry. All populations were also analyzed for the chromosome number. We identified significant differences in the 2C nuclear DNA content among the analyzed species, and this parameter correlated with their ploidy. Diploid (2n = 2x = 24) species N. sibirica and N. tangutorum were found to have smaller genome size (1.24–1.34 and 1.57–1.65 pg) as compared to tetraploid (2n = 4x = 48) species N. komarovii (2.23–2.32 pg), N. pamirica (3.10–3.30 pg), and N. schoberi (2.93–3.39 pg). Intra-population genome size variation was found in examined species, varying from 1.01 to 1.08-fold. Nitraria sibirica has lower inter-population variation of the 2C (1.08-fold) as compared to N. schoberi (1.16-fold). All the Nitraria species are mixoploids. It turned out that an increase in the equatorial axis of Nitraria pollen is associated with an increase in 2C and 1Cx. An exception is N. komarovii, with its intermediate 2C DNA content and the smallest pollen grains. In general, our data confirm Bobrov's previous views on the system of the genus Nitraria, which distinguished ser. Sibiricae and ser. Schoberianae in sect. Nitraria.
... The Syzygium genus belongs to the Myrtaceae family-the eighth largest family of flowering plants-and includes economically important species such as eucalyptus, myrtle, and guava Christenhusz and Byng, 2016;Saber et al., 2023). Although the majority of species of the Myrtaceae family are diploids (2n = 22) with small to intermediate genome sizes (234-1785 Mb), occasional polyploids derived from the most conserved chromosome number x = 11 were also reported (e.g., within the Eugenia, Syzygium, and Psidium genera) (Wilson, 2010;Grattapaglia et al., 2012;Tuler et al., 2019;Pellicer and Leitch, 2020;Machado and Forni-Martins, 2022). The Eucalyptus grandis genome was released in 2014 as the first reference genome for the Myrtales order and the Myrtaceae family (Myburg et al., 2014). ...
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Syzygium is a large and diverse tree genus in the Myrtaceae family. Genome assemblies for clove (Syzygium aromaticum, 370 Mb) and sea apple (Syzygium grande, 405 Mb) provided the first insights into the genomic features and evolution of the Syzygium genus. Here, we present additional de novo chromosome-scale genome assemblies for Syzygium malaccense, Syzygium aqueum, Syzygium jambos, and Syzygium syzygioides. Genome profiling analyses show that S. malaccense, like S. aromaticum and S. grande, is diploid (2n = 2x = 22), while the S. aqueum, S. jambos, and S. syzygioides specimens are autotetraploid (2n = 4x = 44). The genome assemblies of S. malaccense (430 Mb), S. aqueum (392 Mb), S. jambos (426 Mb), and S. syzygioides (431 Mb) are highly complete (BUSCO scores of 98%). Comparative genomics analyses showed conserved organization of the 11 chromosomes with S. aromaticum and S. grande, and revealed species-specific evolutionary dynamics of the long terminal repeat retrotransposon elements belonging to the Gypsy and Copia lineages. This set of Syzygium genomes is a valuable resource for future structural and functional comparative genomic studies on Myrtaceae species.
... Some studies have found that polyploids are more widely distributed than diploids , while others have found no significant differences in the geographic distribution of polyploids and diploids, simply making polyploids at each ploidy level reach their specific ecological niche (Martin and Husband 2009;Liu, et al. 2015). Some studies have found that the ecological niches of polyploids appear separated from diploids, while others have found a large degree of overlap between polyploids and diploids (Thompson, et al. 2014;Arnold, et al. 2015;Hülber, et al. 2015;Liu, et al. 2015;Blaine Marchant, et al. 2016;Sonnleitner, et al. 2016;Visger, et al. 2016;Tuler, et al. 2019). Therefore, we also need more insight into the linkages between WGD, phenotypic variation, and the direct and indirect effects of species interactions when comparing the traits and geographical distribution of polyploids and diploids (Segraves 2017), and cannot generalize. ...
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... Polyploidy has been considered one of the main genomic changes that results in genetic 8 and epigenetic modifications 14 influencing the population genetic structure, ecological niche differentiation, diversification and speciation in plants 15 . About it, Psidium is one outstanding example of the polyploid impact in speciation and geographic distribution 8,15 . Euploidy (autopolyploidy, true allopolyploidy or segmental allopolyploidy) plays a central role in shaping and restructuring plant genomes 15 . ...
... Accesses belonging to P. guajava (2C difference = 0.13 pg among individuals), P. guineense (2C = 0.20 pg) and P. acidum (2C = 0.08 pg) showed less variation of the nuclear 2C value. These nuclear 2C values differences are lower than the 1Cx value of P. guajava (0.475 pg) and P. oblongatum (0.490 pg) determined considering the basic chromosome number (x = 11) of the genus Psidium and the ploidy level of these species (2n = 2x = 22 chromosomes -diploid 8 ). Therefore, the intraspecific variation found among accesses of P. guajava, P. guineense and P. acidum is probably a consequence of secondary metabolites that interfere with the intercalation of the propidium iodide fluorochrome to DNA in the staining step for nuclear suspension preparation for flow cytometry. ...
... IV Cat (5) , Gau (1) , Gau (28) ,Fri (1) , Psi ( (1) , Gua (2) , Gua (1) , Hib (2) , Gui (1) (3) , Cat (1) , Gua (3) , Gua (7) , Gua (18) , Myr (1) , Mac (1) , Psi (2) 37.40 36.72 -38.17 V Cat (12) , Cat (1) , Gua (2) , Gua (8) , Gua (6) , Myr (4) , Myr (11) , Myr (1) , Gau (1) , Gau (1) , Fri (1) , Psi (1) 38.92 38.29 -39.62 VI Aci (2) , Cat (5) , Cat (2) , Gua (1) , Gua (3) , Myr (1) , Myr (7) , Gau (4) , Obl (01) 40.14 39.67 -40.74 VII C at (6) , Cat (2) , Cat (2) , Gui (1) , Myr (2) The GC% values of the evaluated 137 individuals ranged from 34.33% for an individual of P. guineense to 48.95% for an individual of P. myrtoides. The largest intraspecific variations (~ 10%) were in P. myrtoides and P. cattleyanum, which are species that also show nuclear 2C value intraspecific variation. ...
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Diploid and polyploid species derived from the euploid series x = 11 occur in the genus Psidium, as well as intraspecific cytotypes. Euploidy in the genus can alter the gene copy number, resulting in several “omics” variations. We revisited the euploidy, reported genomic (nuclear 2C value, GC%, and copy number of secondary metabolism genes) and epigenomic (5-mC%) differences in Psidium, and related them to essential oil yield and composition. Mean 2C values ranged from 0.90 pg (P. guajava) to 7.40 pg (P. gaudichaudianum). 2C value is intraspecifically varied in P. cattleyanum and P. gaudichaudianum, evidencing cytotypes that can be formed from euploid (non-reduced) and/or aneuploid reproductive cells. GC% ranged from 34.33% (P. guineense) to 48.95% (P. myrtoides), and intraspecific variations occurred even for species without 2C value intraspecific variation. Essential oil yield increased in relation to 2C value and to GC%. We showed that P. guajava (diploid) possesses two and P. guineense (tetraploid) four copies of the one specific TPS gene, as well as eight and sixteen copies respectively of the conserved regions that occur in eight TPS genes. We provide a wide “omics'' characterization of Psidium and show the outcome of the genome and epigenome variation in secondary metabolism.
... Major emphasis has been in Eucalyptus species, for which a reference genome for E. grandis [17] and a multi-species SNP platform EuCHIP60K [18] are available. Alike Eucalyptus, Psidium displays a chromosome number x = 11, a basic complement largely conserved across the family [19,20]. Recently, a chromosome-level assembly of the P. guajava genome corroborated its high collinearity to the E. grandis genome [21]. ...
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Tropical fruit tree species constitute a yet untapped supply of outstanding diversity of taste and nutritional value, barely developed from the genetics standpoint, with scarce or no genomic resources to tackle the challenges arising in modern breeding practice. We generated a de novo genome assembly of the Psidium guajava, the super fruit "apple of the tropics", and successfully transferred 14,268 SNP probesets from Eucalyptus to Psidium at the nucleotide level, to detect genomic loci linked to resistance to the root knot nematode (RKN) Meloidogyne enterolobii derived from the wild relative P. guineense. Significantly associated loci with resistance across alternative analytical frameworks, were detected at two SNPs on chromosome 3 in a pseudo-assembly of Psidium guajava genome built using a syntenic path approach with the Eucalyptus grandis genome to determine the order and orientation of the contigs. The P. guineense-derived resistance response to RKN and disease onset is conceivably triggered by mineral nutrients and phytohormone homeostasis or signaling with the involvement of the miRNA pathway. Hotspots of mapped resistance quantitative trait loci and functional annotation in the same genomic region of Eucalyptus provide further indirect support to our results, highlighting the evolutionary conservation of genomes across genera of Myrtaceae in the adaptation to pathogens. Marker assisted introgression of the resistance loci mapped should accelerate the development of improved guava cultivars and hybrid rootstocks.
... Psidium L. presents polyploid species, with the basic number defined as x=11 (Atchison, 1947;Costa and Forni-Martins, 2006). Polyploidy has played an important role in the evolution and diversification of the genus (Marques et al., 2016;Tuler et al., 2019;Machado et al., 2021), in which records of diploid, triploid, tetraploid, hexaploid, and octaploid species are common (Costa and Forni-Martins, 2006;Tuler et al., 2015Tuler et al., , 2019Machado et al., 2021). The species Psidium cattleyanum Sabine also present a polyploid serie with different chromosome numbers reported, such as 2n=33, 44, 55, 66, 77, 88, 99, 100, 110, and 132 (Atchison, 1947;Costa and Forni-Martins, 2006;Hirano and Nakasone, 1969;Medina, 2014;Souza et al., 2015;Machado et al., 2021Machado et al., , 2022. ...
... Psidium L. presents polyploid species, with the basic number defined as x=11 (Atchison, 1947;Costa and Forni-Martins, 2006). Polyploidy has played an important role in the evolution and diversification of the genus (Marques et al., 2016;Tuler et al., 2019;Machado et al., 2021), in which records of diploid, triploid, tetraploid, hexaploid, and octaploid species are common (Costa and Forni-Martins, 2006;Tuler et al., 2015Tuler et al., , 2019Machado et al., 2021). The species Psidium cattleyanum Sabine also present a polyploid serie with different chromosome numbers reported, such as 2n=33, 44, 55, 66, 77, 88, 99, 100, 110, and 132 (Atchison, 1947;Costa and Forni-Martins, 2006;Hirano and Nakasone, 1969;Medina, 2014;Souza et al., 2015;Machado et al., 2021Machado et al., , 2022. ...
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Polyploidy, a numerical alteration of the karyotype, is one of the most important mechanisms in plant speciation and diversification, but could also be detected among populations, the cytotypes. For example, Psidium cattleyanum, a polyploid complex, has chromosome numbers ranging from 2n=3x=33 to 2n=12x=132. Polyploidization causes an increase in DNA content, and both modifications may cause alteration in plant growth, physiology, and epigenetics. Based on this possibility, here we aim to verify the influence of the polyploidization on the production of P. cattleyanum essential oil chemotypes. Differences in the DNA contents, as a proxy to different ploidies, were observed and three distinct chemotypes were identified through the chromatographic profile analysis. The Psidium cattleyanum DNA content and qualitative and quantitative characteristics of the essential oils presented a positive relationship. Plants with higher DNA contents presented higher levels of oil production, which was mostly composed of hydrogenated sesquiterpenes, while plants with lower DNA contents produced lower amount of oil, which was mostly composed of hydrogenated monoterpenes. Based on the importance of essential oils, polyploid plants, which present higher DNA content, are recommended as possible matrices for the propagation of new plants with the potential to produce major compounds of agronomic and pharmacological interest.
... Intraspecific ploidy variation is reported in many plant groups (Husband et al. 2013), including some species of Psidium L. (Myrtaceae Juss.), as demonstrated by Marques et al. (2016) and Tuler et al. (2019). Psidium has a large number of polyploid records, with chromosome numbers ranging from 2n = 22 to 132, following the basic chromosome number for Myrtaceae, x = 11 (Atchison 1947;Srivastava 1977;Vijayakumar and Subramanian 1985;Costa and Forni-Martins 2006;Souza et al. 2014;Marques et al. 2016;Tuler et al. 2019;Machado et al. 2020Machado et al. , 2022Proença et al. 2022). ...
... Intraspecific ploidy variation is reported in many plant groups (Husband et al. 2013), including some species of Psidium L. (Myrtaceae Juss.), as demonstrated by Marques et al. (2016) and Tuler et al. (2019). Psidium has a large number of polyploid records, with chromosome numbers ranging from 2n = 22 to 132, following the basic chromosome number for Myrtaceae, x = 11 (Atchison 1947;Srivastava 1977;Vijayakumar and Subramanian 1985;Costa and Forni-Martins 2006;Souza et al. 2014;Marques et al. 2016;Tuler et al. 2019;Machado et al. 2020Machado et al. , 2022Proença et al. 2022). Psidium cattleyanum Sabine has eleven cytotypes reported in the literature: 2n = 44, 46, 48, 55, 58, 66, 77, 82, 88,110, and 132 (Atchison 1947;Hirano and Nakazone 1969;Singhal et al. 1985;Raseira and Raseira 1996;Costa and Forni-Martins 2006;Souza et al. 2014;Souza-Pérez and Speroni 2017;Tuler et al. 2019;Machado et al. 2020Machado et al. , 2022Proença et al. 2022). ...
... Psidium has a large number of polyploid records, with chromosome numbers ranging from 2n = 22 to 132, following the basic chromosome number for Myrtaceae, x = 11 (Atchison 1947;Srivastava 1977;Vijayakumar and Subramanian 1985;Costa and Forni-Martins 2006;Souza et al. 2014;Marques et al. 2016;Tuler et al. 2019;Machado et al. 2020Machado et al. , 2022Proença et al. 2022). Psidium cattleyanum Sabine has eleven cytotypes reported in the literature: 2n = 44, 46, 48, 55, 58, 66, 77, 82, 88,110, and 132 (Atchison 1947;Hirano and Nakazone 1969;Singhal et al. 1985;Raseira and Raseira 1996;Costa and Forni-Martins 2006;Souza et al. 2014;Souza-Pérez and Speroni 2017;Tuler et al. 2019;Machado et al. 2020Machado et al. , 2022Proença et al. 2022). Despite the already known number of cytotypes, most of the cytogenetic analysis of this species is based on counting and measuring chromosomes with conventional staining. ...
Article
Polyploidy is a ubiquitous process in angiosperms, arising from autopolyploidy or allopolyploidy. However, few natural polyploid complexes in tropical regions have been well described. Psidium cattleyanum Sabine (Myrtaceae) is a neotropical fruit species with wide distribution and several ploidy levels reported in the literature, being characterized as a polyploid complex. In this study, we provide cytogenetic data of P. cattleyanum based on conventional (chromosome numbers, distribution, and location of CMA/DAPI bands) and molecular techniques (distribution and location of rDNA sites) and flow cytometry (genome size). Moreover, we analyzed the geographic distribution of P. cattleyanum cytotypes. For 75 individuals analyzed in this study, chromosome counts showed nine cytotypes (2n = 3x = 33 to 12x = 132), with two new records (2n = 3x = 33 and 9x = 99). Our results support the occurrence of a polyploid series in P. cattleyanum, derived by the basic number x = 11. We performed CMA/DAPI banding and FISH technique for eight and six cytotypes, respectively. The number of chromosomes, CMA/DAPI bands and 18S and 5S rDNA sites, and DNA content increased linearly related to ploidy level, suggesting the multiplication of the same genome (x = 11) in the different cytotypes. The exception to this gradual increase in DNA content was observed in the cytotypes 2n = 110 and 132, in which the values were relatively lower than expected. There was weak relation between the ploidy and latitudinal distribution. Therefore, Psidium cattleyanum is a good group to study the role of polyploidization events in the evolutionary processes of natural populations, due to the diversity of chromosome numbers and environmental conditions in which cytotypes occur.
... Fonte:CAVALLIN et al., 2016. Devido os caracteres crípticos as espécies desta família botânica possuem muitas semelhanças, principalmente anatômicas, mas de forma geral apresentam tais características:Número de cromossomos, X=11 é conservado nesta família botânica(TULER et al., 2019); os estudos cromossômicos em Myrtaceaes neotropicais (Myrtoideaetem indicação de que a poliploidia é frequente, destoando da característica geral da família Myrtaceae que é a predominância de espécies diploides) ainda são escassos, a maior parte dos estudos realizados são em espécies das subfamílias Chamelaucioideae eLeptospermoideae, incidentes nas regiões da Austrália e no continente Africano (COSTA, Trata-se de um grupo monofilético, que possui em seus aspectos foliares as seguintes características: Epiderme uniestratificada, tricomas unicelulares, folhas hipoestomáticas, estômatos paracíticos, mesófilo dorsiventral, nervura mediana com feixes bicolaterais, cavidades secretoras em ambas as faces e feixes vasculares da nervura central em forma de arco (GOMES, 2009; ASSUNÇÃO; PAULA; PAULA, 2015). Apresentam folhas simples e opostas, com venação broquidódroma, nervura intramarginal, geralmente afastada da margem foliar. ...
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... Due to the difficulty of obtaining direct empirical data, measuring the extent of polyploidy effects can be difficult (Baduel et al. 2018), and knowledge about the influence of whole-genome duplications (WGD) events in the diversification, speciation, and ecology of species remains scarce (Tuler et al. 2019), especially for tropical lineages. For example, autopolyploid organisms are sometimes indistinguishable from their diploid progenitors but evolve as distinct lineages (Baduel et al. 2018;Soltis et al. 2007). ...
... Polyploidy has played an important role in the evolution and diversification of Psidium (Marques et al. 2016;Tuler et al. 2019;Machado et al. 2021), in which records of diploid, triploid, tetraploid, hexaploid, and octaploid species are common (Costa and Forni-Martins 2006;Marques et al. 2016;Tuler et al. 2015Tuler et al. , 2019Machado et al. 2021). Tuler et al. (2019), using SSR, evaluated the relationship between genetic diversity and geographic range in Psidium. ...
... Polyploidy has played an important role in the evolution and diversification of Psidium (Marques et al. 2016;Tuler et al. 2019;Machado et al. 2021), in which records of diploid, triploid, tetraploid, hexaploid, and octaploid species are common (Costa and Forni-Martins 2006;Marques et al. 2016;Tuler et al. 2015Tuler et al. , 2019Machado et al. 2021). Tuler et al. (2019), using SSR, evaluated the relationship between genetic diversity and geographic range in Psidium. ...
Article
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Polyploidy is defined as the presence of more than two complete chromosome sets in an organism and has frequently occurred throughout the history of angiosperms. Polyploidization is a process that typically results in instant speciation. Using Psidium cattleyanum, a natural polyploid complex with several cytotypes, we aim to test two hypotheses regarding speciation in polyploids: polyploidization promotes (1) interruption of gene flow and (2) intraspecific niche divergence. We analyzed 12 natural populations of P. cattleyanum, integrating population genetics data, accessed by microsatellite markers, and climatic niche analysis, using environmental niche modeling, to provide insights about polyploid speciation. We found strong genetic structure in populations and cytotypes and low environmental niche similarity between cytotypes. Genetic diversity declines with increasing ploidy levels which is probably associated with asexual reproduction. Our results corroborate that polyploidy is generating a reproductive barrier and is associated with niche divergence among cytotypes. Therefore, we infer future divergent lineages between cytotypes of P. cattleyanum and confirm the role of polyploidy as an evolutionary step in speciation in this group. Additionally, this study provides new information for the discussion about how polyploidy affects the genetic diversity of taxa and ecological niches.
... The genus Psidium, which belongs to the family Myrtaceae, has a high therapeutic potential for pain and inflammation (Santos et al. 1998;Sekhar et al. 2014;Souza et al. 2019). In Brazil, this genus is widely distributed in the north, northeast, and southeast regions, is prevalent in the Amazon, Atlantic Forest, Cerrado, and Caatinga regions, and has been applied for "in natura" consumption and in agricultural and pharmaceutical industries (Calvi et al. 2017;Tuler et al. 2019). Psidium glaziovianum is a smallto medium-sized tree that reaches up to 5 m in height, with pinnate compound leaves, 4-65 cm leaflets, smooth, dark red to purple oval fruit with sweet acid red pulp, and 5-7 seeds (Landrum 2017). ...
Article
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Plants of the genus Psidium have been employed in “in natura” consumption and agroindustry, and owing to the diversity of phytochemicals, the development of new pharmaceutical forms has received remarkable research interest. In this study, the essential oil obtained from Psidium glaziovianum (PgEO) leaves were evaluated antinociceptive and anti-inflammatory activities were evaluated in mouse models. Initially, PgEO was characterized by gas chromatography–mass spectrometry and gas chromatography with flame ionization detection, and the profile was dominated by sesquiterpene compounds. In the evaluation of acute antinociceptive activity (abdominal contortions induced by acetic acid, formalin, tail immersion, and hot plate tests), PgEO promoted a reduction in nociception in the chemical and thermal models. Additionally, the potential underlying mechanism was investigated using pain pathway blockers, and the results revealed a combined action of opioidergic and muscarinic pathways. The anti-inflammatory potential was confirmed by anti-edematogenic action, reduced cell migration, pro-inflammatory cytokine production, and granuloma formation in chronic processes. This study provides evidence that PgEO can be effective for the treatment of pain and acute and chronic inflammation.