Micrometer insights into Nepeta genus: Pollen micromorphology unveiled.

This study provides a comprehensive pollen micromorphology within the Nepeta genus, revealing intricate details about the pollen grains' structure and characteristics. The findings shed light on the evolutionary and taxonomical aspects of this plant genus, offering valuable insights for botanists and researchers studying Nepeta species. The pollen grains of 18 Nepeta species were studied using scanning electron microscopy (SEM) and light microscopy (LM) in Northern Pakistan. At the microscale, pollen quantitative measurements, qualitative traits, and diverse sculpturing patterns were reported and compared. Significant differences in pollen size, shape, ornamentation, and sculpturing patterns were discovered among the Nepeta species. Our data show that exine sculpturing is quite diverse, with most species exhibiting a reticulate perforate pollen pattern. Nepeta connata, Nepeta discolor, Nepeta elliptica, revealed a distinct bireticulate perforate exine stratification. Hexazonocolpate pollen is the most common. Furthermore, the surface membrane attributes of the colpus varied greatly, ranging from rough, scabrate, psilate, to sinuate patterns. Principal Component Analysis (PCA) was used to discover the key factors influencing pollen diversity. PCA results showed that polar and equatorial diameters, colpi size, and exine thickness were the most influential pollen features between Nepeta species. This study adds to our understanding of pollen morphology in the Nepeta genus, offering information on the vast range of characteristics found in this economically important group. The extensive characterization of pollen features provides useful insights for the categorization and differentiation of Nepeta species, adding to the Lamiaceae micromorphology.

Morphology of pollen grains and orbicules of two threatened species of Cedrela P. Browne (Meliaceae A. Juss.).

Pollen grain morphology and the characterization of additional structures related to pollen dispersion can help to understand the strategies presented by species as well as their taxonomic circumscription. This work investigates the morphology of pollen grains and orbicules of Cedrela fissilis Vell. and Cedrela odorata L., two threatened species of the genus Cedrela P. Browne (Meliaceae A. Juss.) in Brazil. Observations of pollen grains and orbicules of the species were carried out pre- and post-chemical treatment of the samples, under light and scanning electron microscopy, aiming at recognizing structures, detailing morphological characterizations and taking measurements, the last of which were submitted to statistical treatment. The results demonstrate that the pollen grains of the two species are medium in size (measuring between 25 and 50µm), radially symmetrical, isopolar, prolate-spheroidal and 4,5-colporate with subcircular to subquadrangular amb and psilate ornamentation; the orbicules also have a smooth surface and measure about 7 to 10µm. Variation was observed in the size of pollen grains, which presented high coefficient of variation and in the orbicules, as well as in the number of apertures.

Anthropogenic pollen indicators: Global food plants and Latin American human indicators in the pollen record.

Pollen-based evidence of human presence is crucial for reconstructing human history. However, information on the morphology of pollen grains of global food plants and regional pollen-based human indicators is scattered in the literature, leading to the risk of overlooking important evidence of human presence. To address this issue, we first compiled a comprehensive overview of 354 major food plants worldwide, creating a paleoecology-friendly format that includes their family, vernacular name, earliest known use, environmental preference, and geographical region. Moreover, we identified the sources of illustrations of their pollen grains for 209 out of 273 different genera of globally relevant food plants in 10 selected pollen atlases. Secondly, we compiled all human indicators from pollen-based paleoecological literature in Latin America (based on 750 references), providing an overview of 212 single-pollen type indicators and identified 95 crucial combinations of pollen types as "human indices", and their corresponding references. Our review datasets aids in distilling human evidence from numerous fossil pollen records worldwide.

Incomplete partitioning of pollinators by Linum suffruticosum and its coflowering congeners.

PREMISE: Linum suffruticosum shows variations in pollinator fit, pollen pickup, and local pollinators that predict pollen deposition rates. The species often coflowers with other Linum species using the same pollinators. We investigated whether L. suffruticosum trait variation could be explained by local patterns of pollinator sharing and associated evolution to reduce interspecific pollen transfer. METHODS: Pollinator observations were made in different localities (single species, coflowering with other congeners). Floral traits were measured to detect differences across populations and from coflowering species. Reproductive costs were quantified using interspecific hand pollinations and measures of pollen-tube formation, combined with observations of pollen arrival on stigmas and pollen-tube formation after natural pollination in allopatric and sympatric localities. RESULTS: The size and identity of the most important pollinator of L. suffruticosum and whether there was pollinator sharing with coflowering species appeared to explain floral trait variation related to pollinator fit. The morphological overlap of the flowers of L. suffruticosum with those of coflowering species varied, depending on coflowering species identity. A post-pollination incompatibility system maintains reproductive isolation, but conspecific pollen-tube formation was lower after heterospecific pollination. Under natural pollination at sites of coflowering with congeners, conspecific pollen-tube formation was lower than at single-species localities. CONCLUSIONS: Trait variation in L. suffruticosum appears to respond to the most important local pollinator. Locally, incomplete pollinator partitioning might cause interspecific pollination, imposing reproductive costs. These reproductive costs may generate selection on floral traits for reduced morphological overlap with coflowering congeners, leading to the evolution of pollination ecotypes.

Pollen morphological investigation of selected species of family Asteraceae from Pakistan by using light and scanning electron microscopy.

A light and scanning electron microscopy technique was conducted to investigate the palynological features of 19 species belonging to 15 genera of the family Asteraceae. The species under investigation yielded pollen in a range of shapes, including spheroidal, prolate, and subprolate. Trizoncolporate, Tricolporate, and Tetracolporate are the three types of pollen apertures observed in examined species. Except for Gazania rigens, which has reticulate ornamentation under SEM, the exine pattern in all studied species is echinate. The majority of the species had isopolar polarity, whereas some members were apolar and heteropolar. The quantitative parameters that are polar to equatorial diameter, P/E ratio, length of colpi, width of colpi, length of the spine, width of spine, and exine thickness was measured by using light microscopy. The Coreopsis tinctoria had the lowest mean polar 19.75 µm to mean equatorial diameter 18.25 µm while the Silybum marianum had the highest polar 44.7 µm to equatorial 48.2 µm value. The value for colpi length/width was highest in Cirsium arvensis and lowest in C. tinctoria, measuring 9.7/13.2 µm and 2.7/4.7 µm, respectively. The spine length varied from 0.5 µm in Sonchus arvensis to 5.5 µm in Calendula officinalis. Verbesina encelioides had the highest exine thickness measurement (3.3 µm), whereas S. arvensis had the lowest (0.3 µm). The pollen surface of Tagetes erectus, has the highest number (65) whereas the lowest spines number (20) is found in S. arvensis. A taxonomic key based on pollen traits is provided for quick identification of the species. Significant implications for the systematics of the Asteraceae family can be drawn from the reported pollen's quantitative and qualitative data. RESEARCH HIGHLIGHTS: SEM and LM technique was used to study the palynological features of pollen grains. Quantitative and qualitative pollen features of 19 species of family Asteraceae are described. Morphological traits of pollen grains are important for the systematics of family Asteraceae.

Unravelling the palaeobiogeographical history of the living fossil genus Rehderodendron (Styracaceae) with fossil and extant pollen and fruit data.

BACKGROUND: The relict genus Rehderodendron (Styracaceae), the species of which are restricted to mostly warm temperate to tropical climate in East Asia today, is known from fossil fruits and pollen in Europe during warmer periods from the lower Eocene to Pliocene. To infer which extant species are most closely related to the fossils, new data of pollen and fruit morphologiesy of six extant species, and additional new data of fossil pollen and previously described fossil fruits of Rehderodendron, are compared. RESULTS: Both fossil pollen and fruits resemble a morphological mixture of the extant species R. indochinense, R. kwantungense, R. macrocarpum, and R. microcarpum, thus implying that these extant taxa and the fossil European taxa represent an old Eurasian lineage, whereas the pollen and fruit morphology of the extant R. kweichowense and R. truongsonense differ considerably from the fossils and other extant species investigated, and are considered to have evolved independently. CONCLUSIONS: The palaeobiogeographical history of Rehderodendron reveals that its fossil members of the European lineage were most prominent during climatic optima such as the Palaeocene-Eocene Thermal Maximum (PETM), Early Eocene Climate Optimum (EECO) and Middle Miocene Thermal Maximum (MMTM). However, when during the Pliocene the climate changed to colder and less humid conditions, the genus went extinct in Europe but migrated eastwards, most likely in two dispersal events along the Tethys Sea prior to extinction. One of the former most westerly stepping stones is suggested by the refugial occurrence of R. microcarpum in the southeastern Himalaya, whereas R. macrocarpum and R. kwangtungense, the taxa distributed more to the east, might have migrated eastwards already before the Miocene.

Combining genome size and pollen morphology data to study species relationships in the genus Daucus (Apiaceae).

BACKGROUND: The genus Daucus (Apiaceae) comprises about 40 wild species and the cultivated carrot, a crop of great economic and nutritional importance. The rich genetic diversity of wild Daucus species makes them a valuable gene pool for carrot improvement breeding programs. Therefore, it is essential to have good knowledge of the genome structure and relationships among wild Daucus species. To broaden such knowledge, in this research, the nuclear DNA content for 14 Daucus accessions and four closely related species was estimated by flow cytometry and their pollen morphology was analyzed by light and scanning electron microscopy (SEM). RESULTS: The flow cytometric analysis showed a 3.2-fold variation in the mean 2C values among Daucus taxa, ranging from 0.999 (D. carota subsp. sativus) to 3.228 pg (D. littoralis). Among the outgroup species, the mean 2C values were 1.775-2.882 pg. The pollen grains of Daucus were tricolporate, mainly prolate or perprolate (rarely) in shape, and mainly medium or small (rarely) in size (21.19-40.38 µm), whereas the outgroup species had tricolporate, perprolate-shaped, and medium-sized (26.01-49.86 µm) pollen grains. In the studied taxa, SEM analysis revealed that exine ornamentation was striate, rugulate, perforate, or the ornamentation pattern was mixed. At the time of shedding, all pollen grains were three-celled, as evidenced by DAPI staining. We also found high positive correlations between the length of the polar axis (P) and the length of the equatorial diameter (E) of pollen grains, as well as between P and P/E. However, when comparing cytogenetic information with palynological data, no significant correlations were observed. CONCLUSIONS: This study complements the information on the nuclear DNA content in Daucus and provides comprehensive knowledge of the pollen morphology of its taxa. These findings may be important in elucidating the taxonomic relationships among Daucus species and can help in the correct identification of gene bank accessions. In a broader view, they could also be meaningful for the interpretation of evolutionary trends in the genus.

The comparative palynomorphological studies on Turkish Muscari genera and close relatives (Asparagaceae-scilloideae) with their taxonomic implications.

By this study, the palynomorphological traits of 49 taxa (53 plant samples) of the genus Muscari which has been the main research topic and also its relatives, Pseudomuscari, Hyacinthella and Bellevalia have been studied in details of both using light and scanning electron microscope. In the end of palynologic works, it was suggested the descriptive or symbolized main pollen types to make easier the separation or identification of the species of Muscari and its relative groups. According to this symbolization, the pollen types of species may be changeable as D-shaped, elliptical, triangular, and spindle shaped and all types could be seen in different members of the studied taxonomic groups in this paper. One of the most important findings of the study is that the taxa of the genus Muscari, Hyacinthella and Bellevalia do not show a main difference in terms of pollen structures and especially they exhibit harmony. Therefore, it does not possible to mention about the special pollen shape own to one species but it is possible to see common pollen types for closely related taxa or taxon groups. As a result, it could be commented that there is not evolved the close taxonomical groups within Asparagaceae in terms of pollen shape and morphologies and there is considerably a high suitability in point of theirs germ cells or microspores among them. RESEARCH HIGHLIGHTS: A detailed palynological description is provided for Muscari and close relative genus using scanning electron microscopy (SEM) and light microscopy (LM). Pollen size and sulcus length are been determined as the best informative pollen characters to understand Muscari and other studied species.

Taxonomic significance of palynological studies for identification of two morphologically similar Malva species.

Pollen studies can assist in distinguishing different plant taxa on the basis of pollen diameter, exine thickness, spines length, spine shape, number of rows of spines between colpi, shape of pollen, P/E ratio, pollen class and aperture type. Light microscopy (LM) and scanning electron microscopy (SEM) was used to compare and differentiate two Malva species on the basis of different pollen characteristics. Pollen in Malva neglecta and Malva parviflora had punctate and subpsilate, and, granulate and verrucate sculpturing between spines respectively. Both species pollen had bulbous spine base, but were more swollen in M. parviflora. M. neglecta had more pollen size, spine length and distance between spine base and spine apex, except spine base and pore diameter. The scatterplot matrix showed a wide variation among sculpturing types and other pollen attributes of both species. In PCA biplot, pore diameter and spine length were linked to M. parviflora while other pollen attributes were clearly linked to M. neglecta, indicating the taxonomic significance of the observed characters. It was concluded that the palynological studies can act as an identification tool and are of great significance in delimiting Malva and different plant taxa. RESEARCH HIGHLIGHTS: Taxonomic identifications is a big problem in morphologically identical Malva neglecta and Malva parviflora. The usefulness of pollen morphology (pollen diameter and exine thickness) and wall ornamentation in identification through light microscopy (LM) and scanning electron microscopy (SEM) was confirmed. Pollen of M. neglecta can be differentiated from M. parviflora by its sculpturing patterns. Spine base and pore diameter of M. parviflora is a taxonomically important feature.

Pollen morphology of selected melliferous plants and its taxonomic implications using microscopy.

This research aimed to explore the melliferous bee flora pollen from arid zones of District Attock, Islamabad Capital and Rawalpindi. Morpho-palynological features has provided key information for the classification and help to explain the taxonomy of several genera and species. Therefore, in this work, we examined the morphological characters of pollen potentially to be used for the species delimitation of bee flora. A total of 18 bee floral species, classified into 12 different families were collected, pressed, identified and then acetolyized to observe the grains under microscopy. Bee pollen can be described by small, medium to large sized morphotypes presenting five types of shapes, prolate spheroidal in (seven species), sub-prolate (six species), oblate spheroidal (three species), oblate and prolate (one species each) was examined. The observed polar and equatorial diameter were found maximum in Zea mays 73.5 µm and Pelargonium inquinans 66.75 µm respectively. Pollen of six different types was found namely: tricolporate, 3-zonocolporate, monoportae, tetracolporate, tetraporate and mononsulcate. Whereas highest value of colpi length was measured in Grewia tenax (24.55 µm). Exine surface nature of pollen was examined echinate psilate and scabrate. Exine thickness noted to be maximum in Verbena tenuisecta (8.40 µm) and minimum in Citrus macrocarpa (0.4 µm). Bee floral species considered difficult to identify based on other morphological traits were successfully distinguished using pollen quantitative and qualitative traits, confirming the importance of pollen morphology to diagnose characters to correctly identify honeybee flora. RESEARCH HIGHLIGHTS: Pollen diversity among honeybee flora. Highly variations observed via microscopy in pollen micromorphology. Palyno-morph has taxonomic value for the identification of honeybee floral species.

Systematic palynology in Korean Piperales with special focus on its exine surface ornamentation and orbicule morphology.

The pollen and orbicule morphology of the Korean Piperales (Aristolochia, Asarum, Houttuynia, Piper, and Saururus) were investigated via scanning electron microscopy. Piperales pollen is a monad, its size ranging from very small to large (P = 7.78-51.4 µm, E = 6.68-43.1 µm), and having a mainly circular to sub-circular shape. The aperture type is constant in the genus [inaperturate (Aristolochia), tri to pentaporate (Asarum), and monosulcate (Houttuynia, Piper, and Saururus)]. There are four distinct types of exine ornamentation: Fossulate with perforate, microreticulate with gemmae, microperforate with granula, and microechinate. The orbicules (minute sporopollenin granules) were observed in all studied taxa and thus, may be a possible symplesiomorphic characteristic of Piperales. Further, the observed orbicule surface ornamentation was similar to pollen exine patterns, for example muri, gemmae, or granula. This resemblance between orbicule and pollen exine ornamentation may imply a similar biosynthesis pattern of sporopollenin of pollen exine and orbicules. The phenogram resulting from a cluster analysis using palynological characters was generally consistent with the known molecular phylogeny of Piperales. This initial study will help understand the palynological diversity and provide detailed information of pollen and orbicule characteristics in Piperales.

Why does pollen morphology vary? Evolutionary dynamics and morphospace occupation in the largest angiosperm order (Asterales).

Morphological diversity (disparity) is a key component of biodiversity and increasingly a focus of botanical research. Despite the wide range of morphologies represented by pollen grains, to date there are few studies focused on the controls on pollen disparity and morphospace occupation, and fewer still considering these parameters in a phylogenetic framework. Here, we analyse morphospace occupation, disparity and rates of morphological evolution in Asterales pollen, in a phylogenetic context. We use a dataset comprising 113 taxa from across the Asterales phylogeny, with pollen morphology described using 28 discrete characters. The Asterales pollen morphospace is phylogenetically structured around groups of related taxa, consistent with punctuated bursts of morphological evolution at key points in the Asterales phylogeny. There is no substantial difference in disparity among these groups of taxa, despite large differences in species richness and biogeographic range. There is also mixed evidence for whole-genome duplication as a driver of Asterales pollen morphological evolution. Our results highlight the importance of evolutionary history for structuring pollen morphospace. Our study is consistent with others that have shown a decoupling of biodiversity parameters, and reinforces the need to focus on disparity as a key botanical metric in its own right.

Southeast Asian Dipterocarp origin and diversification driven by Africa-India floristic interchange.

The evolution and diversification of ancient megathermal angiosperm lineages with Africa-India origins in Asian tropical forests is poorly understood because of the lack of reliable fossils. Our palaeobiogeographical analysis of pollen fossils from Africa and India combined with molecular data and fossil amber records suggest a tropical-African origin of Dipterocarpaceae during the mid-Cretaceous and its dispersal to India during the Late Maastrichtian and Paleocene, leading to range expansion of aseasonal dipterocarps on the Indian Plate. The India-Asia collision further facilitated the dispersal of dipterocarps from India to similar climatic zones in Southeast Asia, which supports their out-of-India migration. The dispersal pathway suggested for Dipterocarpaceae may provide a framework for an alternative biogeographic hypothesis for several megathermal angiosperm families that are presently widely distributed in Southeast Asia.

Palyno-morphological attributes of some selected plant species of family Asteraceae from district Dera Ismail Khan, KPK, Pakistan.

The current study aimed to investigate the palyno-morphological features of Asteraceous species from District Dera Ismail Khan, Khyber Pakhtunkhwa, Pakistan. A total of 13 species were collected, pressed, identified, and examined by using light and scanning electron microscopy. Both qualitative and quantitative pollen characters were observed, that is, equatorial and polar diameter, number of pores and colpi, exine thickness, exine sculpturing, pollen shape, and P/E ratio. Pollen shapes observed in studied taxa were spheroidal, suboblate, oblate-spheroidal, and lophate-spheroidal. The aperturation patterns of pollens vary from tricolporate to tetracolporate. The maximum polar and equatorial diameter of 75-100 (87.5 ± 13.6) µm and 87.5-117.5 (102.5 ± 16.4) µm, respectively, was recorded in Echinops echinatus, while minimum polar and equatorial diameter of 20-22.5 (21 ± 1.36) µm and 22.5-25 (23.5 ± 1.36) µm, respectively, was recorded in Aster subulatus. Six types of exine sculpturing patterns were observed; echinate, micro-echinate, echinate-microreticulate, microreticulate, echinate-perforate, and scabrate. Maximum exine thickness of 8.75 µm was recorded in E. echinatus and minimum of 1.25 µm in Launaea mucronata. The pollen morphology has valuable significance in understanding the taxonomy of different plant groups and plays a central role in the correct identification and classification of Asteraceous flora at species, generic, and tribe levels.

Pollen morphology of the genus Sophora (Fabaceae) and its taxonomic implications.

The genus Sophora (Fabaceae) is one of the taxonomically challenging genera with high economic and medical values. In this study, the pollen morphology of 43 samples of 27 species, 4 subspecies, and 4 varieties of the genus Sophora and 3 closely related genera was examined using scanning electron microscopy to evaluate the pollen diversity of the genus and its taxonomic significance. Pollen grains of the studied species were tricolporate (rarely six-aperture), and pollen shape varied from suboblate, spheroidal, subprolate to prolate. Echinate external ornamentation was reported for the first time in some species of the genus. Aperture membrane ornamentation and outline in a polar/equatorial view were described for the first time in the genus. Principal component analysis was used to understand the relationship and discrimination between the species and the genera, with six components accounting for 79.92% of the total variance. Taxonomic keys based on pollen morphology were also constructed to easily identify the taxa of the genus through palynological characteristics. Results showed that pollen morphology alone is not sufficient to elucidate or reconstruct taxonomic relationships within the genus Sophora, but palynological assessments can provide some useful information for identifying taxonomically problematic taxa.

Pollen morphology and its taxonomic potential in some selected taxa of Caesalpiniaceae observed under light microscopy and scanning electron microscopy.

Pollen micromorphology is not only used to check the functional and structural evolution in plants but also to solve the taxonomic problem related to the classification of plants. Therefore, keeping in view the significance of pollen traits, selected taxa of the subfamily Caesalpiniaceae was collected from different geographical regions of Pakistan. The species were then analyzed under both light microscopy and scanning electron microscopy techniques to investigate the importance of micromorphological characters of pollen in the identification and classification of species. Great variation was recorded in equatorial shape, surface ornamentation, tectum, polar diameter, equatorial diameter, and exine thickness. However, little variation was observed in pollen type, polar shape, and fertility of pollen. The equatorial shape of five types was observed: prolate, prolate-spheroidal, spheroidal-subprolate, subspheroidal-prolate, and subspheroidal. Four types of surface ornamentation, psilate, granulate, clavate, and perforate, were recorded. Tectum of five types, intactate, reticulate regulate, medium reticulate, tactate, and striate, was observed. Sexine was thicker than nexine in all studied species. The largest polar diameter was observed in Caesalpinia pulcherrima 64.1 µm while the smallest in Parkinsonia aculeata 26.1 µm. The largest equatorial diameter was found in C. pulcherrima of 70.25 µm whereas the smallest in P. aculeata 27.57 µm. All the pollens analyzed were tricolporate. All studied species have a fertility ratio of more than 90%. A taxonomic key was developed to show the variation in pollen features and delimit species for the correct identification. In conclusion, the pollen traits were found useful to define species boundaries at various taxonomic ranks and will strengthen the taxonomy of this subfamily. Besides, this study also explored the palynological traits and their implication in the taxonomy of the subfamily Caesalpiniaceae.

Morpho-palynological investigation of gymnospermal flora from subalpine and alpine zones of northern Pakistan using LM and SEM.

The pollen morphology, with special reference to exine sculpture, of some species of the gymnosperms was assessed for the first time from the subalpine and alpine zones of western Himalayas northern Pakistan. The pollen of all these species is airborne and allergenic, so pollen morphology helps for identification of this allergenic pollen at specific level. Different morpho-palynological characteristics were analyzed including size range of pollen, polar and equatorial diameter ratio, exine ornamentation, sculpturing, exine thickness, pollen type, and shape. For accurate and quick identification of species, taxonomic key was made based on different morpho-palynological characteristics. The quantitative data were processed using SPSS software. Gymnospermal pollen includes inaperturate, rarely 1-colpate observed in (Cupressaceae), hexazonocolpate in (Ephedraceae), vesiculate, bissacate in (Pinaceae), and inaperturate in (Taxaceae). Different pollen shapes observed were prolate (4 spp), sub-spheroidal (7 spp), and oblate (1 spp). Variation was observed in exine sculpturing granular (4 spp), reticulate (1 spp), areolate-punctate (3 spp), and psilate (2 spp). This is based on the analysis of 10 plants belonging to four families of gymnosperms. Distinct pollen shape has emerged as the most diagnostic feature to separate some genera such as spheroidal in (Cupressaceae, Taxaceae), prolate and radiosymmetrical in (Ephedraceae), and bilateral in (Pinaceae). Exine thickness and sculpturing proved to be helpful at generic and specific levels. The results reinforced the significance of gymnospermal pollen morphological features which were used as aid for valuable taxonomic tool in plant systematics.

Morphological differences between anemophilous and entomophilous pollen.

In order to provide a palynological guide for the identification of insect-carrying pollen grains, we studied the pollen grains of 10 anemophilous species and 10 entomophilous species in the Beijing urban area using light and scanning electron microscopies. We found that anemophilous pollen grains are small, spheroidal, or oblate spheroidal, while entomophilous pollen grains are medium and oblate. Comparison of the exine thickness and surface ornamentation showed that anemophilous pollen grains have significantly thinner exine and smoother surface ornamentation than entomophilous pollen grains. The results also revealed pollen characteristics adaptive to different pollination types. Overall, our study indicated that pollen morphology might be helpful for preliminary identification of anemophilous and entomophilous pollen.

A Comprehensive Study of the Genus Sanguisorba (Rosaceae) Based on the Floral Micromorphology, Palynology, and Plastome Analysis.

Sanguisorba, commonly known as burnet, is a genus in the family Rosaceae native to the temperate regions of the Northern hemisphere. Five of its thirty species are distributed in Korea: Sanguisorba officinalis, S. stipulata, S. hakusanensis, S. longifolia, and S. tenuifolia. S. officinalis has been designated as a medicinal remedy in the Chinese and Korean Herbal Pharmacopeias. Despite being a valuable medicinal resource, the morphological and genomic information, as well as the genetic characteristics of Sanguisorba, are still elusive. Therefore, we carried out the first comprehensive study on the floral micromorphology, palynology, and complete chloroplast (cp) genome of the Sanguisorba species. The outer sepal waxes and hypanthium characters showed diagnostic value, despite a similar floral micromorphology across different species. All the studied Sanguisorba pollen were small to medium, oblate to prolate-spheroidal, and their exine ornamentation was microechinate. The orbicules, which are possibly synapomorphic, were consistently absent in this genus. Additionally, the cp genomes of S. officinalis, S. stipulata, and S. hakusanensis have been completely sequenced. The comparative analysis of the reported Sanguisorba cp genomes revealed local divergence regions. The nucleotide diversity of trnH-psbA and rps2-rpoC2, referred to as hotspot regions, revealed the highest pi values in six Sanguisorba. The ndhG indicated positive selection pressures as a species-specific variation in S. filiformis. The S. stipulata and S. tenuifolia species had psbK genes at the selected pressures. We developed new DNA barcodes that distinguish the typical S. officinalis and S. officinalis var. longifolia, important herbal medicinal plants, from other similar Sanguisorba species with species-specific distinctive markers. The phylogenetic trees showed the positions of the reported Sanguisorba species; S. officinalis, S. tenuifolia, and S. stipulata showed the nearest genetic distance. The results of our comprehensive study on micromorphology, pollen chemistry, cp genome analysis, and the development of species identification markers can provide valuable information for future studies on S. officinalis, including those highlighting it as an important medicinal resource.

A species-specific functional module controls formation of pollen apertures.

Pollen apertures are an interesting model for the formation of specialized plasma-membrane domains. The plant-specific protein INP1 serves as a key aperture factor in such distantly related species as Arabidopsis, rice and maize. Although INP1 orthologues probably play similar roles throughout flowering plants, they show substantial sequence divergence and often cannot substitute for each other, suggesting that INP1 might require species-specific partners. Here, we present a new aperture factor, INP2, which satisfies the criteria for being a species-specific partner for INP1. Both INP proteins display similar structural features, including the plant-specific DOG1 domain, similar patterns of expression and mutant phenotypes, as well as signs of co-evolution. These proteins interact with each other in a species-specific manner and can restore apertures in a heterologous system when both are expressed but not when expressed individually. Our findings suggest that the INP proteins form a species-specific functional module that underlies formation of pollen apertures.