Authorship：Lead author   Language：Japanese   Publishing type：Research paper (bulletin of university, research institution)  

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Language：English   Publishing type：Research paper (scientific journal)   Publisher：Wiley  

DOI： 10.1002/jez.2408

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Other Link： https://onlinelibrary.wiley.com/doi/full-xml/10.1002/jez.2408

Authorship：Lead author   Language：Japanese   Publishing type：Research paper (scientific journal)  

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The Japanese wrinkled frog Glandirana rugosa is separated into five genetically different groups. One group in western Japan is further divided into three subgroups, found in Kyushu, Shikoku, and western Honshu. We collected G. rugosa frogs at 39 sites in Kyushu and determined nucleotide sequences of the mitochondrial 12S and 16S rRNA genes for phylogenetic analysis. Unexpectedly, we found a group of frogs in southeastern Kyushu that did not cluster with any of the pre-existing five groups of G. rugosa on the phylogenetic trees. The frogs in the new group and G. rugosa in Kyushu were externally similar, but there were a few significant differences in morphological features between the two populations. In addition, we observed significant differences in the frogs' calls . Thus, the group of the frogs in southeastern Kyushu may represent a new candidate species in the genus Glandirana. We discuss the possibility of a new species.

DOI： 10.2108/zs190007

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Language：English   Publishing type：Research paper (scientific journal)  

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Language：Japanese   Publishing type：Research paper (scientific journal)   Publisher：愛媛大学教育・学生支援機構  

CiNii Books

CiNii Research

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Other Link： http://id.ndl.go.jp/bib/029914590

Language：Japanese   Publishing type：Research paper (scientific journal)  

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Language：Japanese   Publishing type：Research paper (scientific journal)   Publisher：愛媛大学教育学部  

CiNii Books

CiNii Research

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Language：English   Publishing type：Research paper (scientific journal)  

In the teleost fish medaka, an adult ovary simultaneously contains developing oocytes at all phases of oogenesis during the breeding season. However, it remains unclear where oocytes at each developmental stage are located in the ovary by the time of ovulation. To examine the relationship between the developmental stage of oocytes and their positions in the ovary of vertebrate medaka during oogenesis, the stage of oocyte development was determined from the diameter of the oocytes and the cellular morphological characteristics, such as the germinal vesicle and micropyle at the animal pole and attaching filaments at the vegetal pole, and the positions of developing oocytes in the ovary in all sections were observed. Furthermore, to investigate the characteristics of the dorsal ovarian epithelium to which the oocytes attach themselves, the dorsal and vegetal ovarian epithelia were observed. The dorsal ovarian epithelium invaginated in spots. When all serial sections of the oocytes were observed, all oocytes at stages I-VIII were attached to the dorsal ovarian epithelium, regardless of whether it invaginated or not.

DOI： 10.2108/zs170210

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Language：Japanese   Publishing type：Research paper (scientific journal)  

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Language：English   Publishing type：Research paper (scientific journal)   Publisher：WILEY  

Each vertebrate species, as a general rule, has either the XX/XY or ZZ/ZW chromosomes by which sex is determined. However, the Japanese Rana (R.) rugosa frog is an exception, possessing both sex-determining combinations within one species, varying with region of origin. We collected R. rugosa frogs from 104 sites around Japan and South Korea and determined the nucleotide sequences of the mitochondrial 12S ribosomal RNA gene. Based on the sequences, R. rugosa frogs were divided into four groups from Japan and one from South Korea. The ZZ/ZW type is reportedly derived from the XX/XY type, although recently a new ZZ/ZW type of R. rugosa was reported. However, it still remains unclear from where the sex chromosomes in the five groups of this species were derived. In this study, we successfully isolated a sex-linked DNA maker and used it to classify R. rugosa frogs into several groupings. From the DNA marker as well as from nucleotide analysis of the promoter region of the androgen receptor (AR) gene, we identified another female heterogametic group, designated, West-Central. The sex chromosomes in the West-Central originated from the West and Central groups. The results indicate that a sex-linked DNA marker is a verifiable tool to determine the origin of the sex chromosomes in R. rugosa frogs in which the sex-determining system has changed, during two independent events, from the male to female heterogamety.

DOI： 10.1002/jez.2130

Web of Science

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Language：Japanese   Publishing type：Research paper (scientific journal)  

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Language：Japanese  

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Language：Japanese   Publishing type：Research paper (scientific journal)  

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Language：English   Publishing type：Research paper (scientific journal)   Publisher：PUBLIC LIBRARY SCIENCE  

INTRODUCTION: In the Japanese frog Rana (R.) rugosa the androgen receptor (AR) gene on the W chromosome (W-AR) is barely expressed. Previously we showed that incomplete female-to-male sex-reversal occurred in Z-AR transgenic female frogs. To date, however, there is no report showing that AR with androgens can determine genetically programed male sex fate in any vertebrate species. Here, we examined whether AR together with androgens functions as a sex determinant in an amphibian species. METHODS: To examine whether complete female-to-male sex-reversal occurs in R. rugosa frogs, we produced AR-transgenic (Tg) and -knockdown (KD) female R. rugosa frogs by the I-SceI meganuclease-mediated gene trap and CRISPR/Cas9 system, respectively. AR-Tg and -KD tadpoles were reared in water containing testosterone (T) at 0 to 7.1 ng/ml. Frozen sections were prepared from the gonads of metamorphosed frogs and immunostained for laminin, Vasa, Pat1a, CYP17 and AR. We also employed PCR analysis to examine Dmrt1, Pat1a and CYP17 expression in the gonads of KD and placebo-KD female frogs. RESULTS: Complete female-to-male sex-reversal occurred in the AR-Tg ZW female frogs when a low dosage of T was supplied in the rearing water of tadpoles. However, no sex-reversal was observed in AR-KD ZW female frogs when the gonads were treated with dosages of T high enough to induce complete female-to-male sex-reversal even in wild type frogs. DISCUSSION: These results suggest that AR with its androgen ligand functions as a male sex-determinant in the ZW type R. rugosa frogs.

DOI： 10.1371/journal.pone.0178067

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Language：Japanese   Publishing type：Research paper (scientific journal)  

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Language：English   Publishing type：Research paper (scientific journal)  

Androgens play a critical role in testicular differentiation in many species of vertebrates. While female-to-male sex reversal can be induced by testosterone (T) in some species of amphibians, the mechanism still remains largely unknown even at the histological level. In this study, we determined a threshold dosage of T to induce female-to-male sex reversal in the Japanese frog Rana (R.) rugosa. Tadpoles were allowed to metamorphose into frogs with T present in the rearing water. At 0.2 ng/mL T, female frogs formed tissue comprising a mixture of ovary and testis, the so-called ovotestis, the size of which was significantly smaller than the wild-type ovary. Histological changes occurring in the oocytes of T-treated ovaries induced oocyte degeneration in the masculinizing ovaries leading to their final disappearance. In parallel, many germ cells emerged in the cortex of the ovotestis and, later, in the medulla as well. RT-PCR analysis revealed upregulated expression of CYP17 and Dmrt1 but not 17βHSD in the ovotestis, and downregulation of Pat1a expression. Furthermore, immunohistology revealed CYP17-positive signals in the cortex of the masculinizing ovary, spreading throughout the whole area as the testis developed. These results indicate that oocytes are sensitive to T in the ovary of R. rugosa and that male-type germ cells expand in the masculinizing gonad (testis) contemporaneous with oocyte disappearance.

DOI： 10.1002/jez.2037

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Language：Japanese   Publishing type：Research paper (bulletin of university, research institution)  

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Language：English   Publishing type：Research paper (scientific journal)  

The phenotypic sex of many species of amphibians is subject to reversal by steroid hormones. The mechanism of this process, however, still remains largely unknown. As a step toward understanding the histological changes during sex reversal in amphibians, we analyzed two- and three-dimensional (2D and 3D) structures of sex-reversing gonads in Rana rugosa frogs. 2D views revealed that many oocytes in the wild-type ovary disappeared during female-to-male sex-reversal concomitant with the emergence of Vasa-positive small germ cells. Some of the germ cells were labeled with BrdU. BrdU-positive germ cells were few in the testosterone (T) treated ovaries at days 8 and 16, which resembled wild-type ovaries. Basement membranes became disrupted by day 24 in T-treated ovaries. However, the membranes were later reconfigured into testis-like gonadal structures 40 days after T treatment. 3D imaging of the sex-reversing gonad using serial immunostained sections showed that germ cells were organized in linear fashion extending out from where the sex-reversing gonad attached to the mesorchium 24 days after T treatment. Germ cells were increased in number by 40 days and were localized to the cortex of the gonads. In a T-untreated testis at day 24, many germ cells were distributed throughout the cortex except in the central space, while the efferent duct ran between two sheets of the mesorchium. These results, taken together, suggest that the mesorchium plays an important role in the organization of testicular structure. This is the first report showing germ cell ontogeny and organization in the female-to-male sex-reversing gonad in a vertebrate species.

DOI： 10.1002/jez.2009

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Language：Japanese   Publishing type：Research paper (scientific journal)  

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Language：Japanese   Publishing type：Research paper (scientific journal)  

CiNii Books

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Other Link： http://iyokan.lib.ehime-u.ac.jp/dspace/handle/iyokan/4697

Language：English   Publishing type：Research paper (scientific journal)  

The Pat1 gene is expressed in the immature oocytes of Xenopus, and is reportedly involved in regulating the translation of maternal mRNAs required for oocyte-maturation. However, it is still unknown when Pat1a first appears in the differentiating ovary of amphibians. To address this issue, we isolated the full-length Pat1a cDNA from the frog Rana rugosa and examined its expression in the differentiating ovary of this frog. Among eight different tissues examined, the Pat1a mRNA was detectable in only the ovary. When frozen sections from the ovaries of tadpoles at various stages of development were immunostained for Vasa-a germ cell-specific protein-and Pat1a, Vasa-immunopositive signals were observed in all of the germ cells, whereas Pat1a signals were confined to the growing oocytes (50-200 μm in diameter), and absent from small germ cells (<50 μm in diameter). Forty days after testosterone injection into tadpoles to induce female-to-male sex-reversal, Pat1a-immunoreactive oocytes had disappeared completely from the sex-reversed gonad, but Vasa-positive small germ cells persisted. Thus, Pat1a would be a good marker for identifying the sexual status of the sex-reversing gonad in amphibians. In addition, fluorescence in situ hybridization analysis showed Pat1a to have an autosomal locus, suggesting that Pat1a transcription is probably regulated by a tissue-specific transcription factor in R. rugosa.

DOI： 10.1002/jez.1938

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Language：Japanese   Publishing type：Research paper (scientific journal)  

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The role of anti-Müllerian hormone (AMH) during gonad development has been studied extensively in many species of mammal, bird, reptile, and fish but remains unresolved in amphibians. In male mammalian embryos, Sox9 activates AMH expression, which initiates regression of the Müllerian ducts. However, Sox9 (Sry-related HMG box 9) is unlikely to initiate AMH in chicken, because AMH precedes Sox9 expression in this species. To clarify whether AMH is involved in testicular differentiation in amphibians, we cloned the full-length AMH cDNA from the Japanese wrinkled frog, Rana rugosa. The AMH gene, which appears to be autosomal, is exclusively expressed in the testis of adult frog among 8 different tissues examined; Sertoli cells are probably responsible for its expression. AMH expression was found in the undifferentiated gonad of both male and female tadpoles, increasing in the differentiating testis. Moreover, we observed consensus binding sites for Sox9 in the 5'-flanking region of the AMH gene. Sox9 stimulated statistically significant AMH expression in luciferase reporter assays when coexpressed in Xenopus kidney-derived A6 cells. However, Sox9 expression showed no sexual dimorphism when AMH expression was up-regulated in the developing testis. These results, taken together, suggest that AMH is probably involved in testicular differentiation in R. rugosa, although an additional, perhaps tissue-specific, transcription factor may be required for the regulation of AMH transcription.

DOI： 10.1210/en.2013-2053

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Language：Japanese   Publishing type：Research paper (scientific journal)  

CiNii Books

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Other Link： https://opac1.lib.ehime-u.ac.jp/iyokan/TD00003251

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CiNii Books

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In vertebrates, gonadal production of steroid hormones is regulated by follicle-stimulating hormone (FSH) and luteinizing hormone (LH) via their receptors designated FSHR and LHR, respectively. We have shown recently that steroid hormones are synthesized in the differentiating gonad of tadpoles during sex determination in the frog Rana rugosa. To elucidate the role of gonadotropins (GTHs) and their receptors in the production of gonadal steroid hormones during sex determination, we isolated the full-length FSHβ, LHβ, FSHR and LHR cDNAs from R. rugosa and determined gonadal expression of FSHR (FSH receptor) and LHR (LH receptor) as well as brain expression of FSHβ and LHβ during sex determination in this species. The molecular structures of these four glycoproteins are conserved among different classes of vertebrates. FSHβ expression was observed at similar levels in the whole brain (including the pituitary) of tadpoles, but it showed no sexual dimorphism during gonadal sex determination. By contrast, LHβ mRNA was undetectable in the whole brain of tadpoles. FSHβ-immunopositive cells were observed in the pituitary of female tadpoles with a differentiating gonad. Furthermore, FSHR expression was significantly higher in the gonad of female tadpoles during sex determination than in that of males, whereas LHR was expressed at similar levels in males and females. The results collectively suggest that FSHR, probably in conjunction with FSH, is involved in the steroid-hormone production during female-sex determination in R. rugosa.

DOI： 10.1016/j.ygcen.2011.04.011

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In certain species of amphibians gonadal differentiation is influenced by steroid hormones. In the case of the frog Rana rugosa testosterone given to tadpoles reverses sex from female to male, while the opposite reversal - male to female - can be achieved using estradiol-17β. In this study, we investigated whether CYP19 (P450 aromatase), the enzyme responsible for a production of estradiol-17β, was present in the differentiating gonad of R. rugosa. Initially, we immunized rabbits against frog CYP19 peptides and performed immunostaining using specific antibodies purified from that serum. CYP19-reactive signals were observed in gonadal somatic cells of the female, but not male tadpoles at stage (St.) I (the stage prior to phenotypic sex determination in tadpoles of R. rugosa). Immunopositive signals were also produced in ovarian somatic cells froglets at St. XXV (just after the completion of metamorphosis). We also examined the enzymatic activity of CYP19 in the differentiating gonad of R. rugosa. Reverse-phase HPLC (high performance liquid chromatography) analysis revealed that [(3)H]testosterone was converted to [(3)H]estradiol-17β in the gonad of tadpoles at St. I. Interestingly, the rate of conversion was much higher in females than in males. To the best of our knowledge, this is the first report on the biosynthesis of estradiol-17β in the gonad of amphibians, and the co-incident identification of active CYP19 enzyme in the differentiating gonad of R. rugosa. Based on our results, we conclude that estradiol-17β may be involved in ovarian differentiation in this species.

DOI： 10.1016/j.ygcen.2010.10.015

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We cloned a cDNA encoding Vasa, a member of the DEAD (Asp-Glu-Ala-Asp) family of proteins, from the ovary of the frog Rana rugosa. Comparative alignment of amino acid sequences with known Vasa from several species of vertebrate showed that the R. rugosa orthologue shares eight conserved regions with Vasa from other vertebrates. Vasa gene expression was restricted to the testis and ovary among ten different tissues examined. Vasa expression showed no sexual dimorphism during sex determination in R. rugosa, but became higher in the ovary thereafter. By Western blot analysis, a single Vasa band with a molecular weight of 80.9 kDa was detected. The same antibody immunohistochemically detected Vasa in a few cells in the embryonic endoderm at stage 15; the beginning of closure of neural folds, and in the cytoplasm of spermatogonia in the testis, and oocytes in the ovary of tadpoles at stage XX; marked by one or both forelegs protruded. Together, these results suggest that Vasa is a highly specific marker of germ cells and hence useful for studies of germ cell specification and function in amphibians as it already is in other species of both invertebrates and vertebrates such as Drosophila and zebrafish.

DOI： 10.1002/jez.617

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The translational regulation of maternal mRNAs is one of the most important steps in the control of temporal-spatial gene expression during oocyte maturation and early embryogenesis in various species. Recently, it has become clear that protein components of mRNPs play essential roles in the translational regulation of maternal mRNAs. In the present study, we investigated the function of P100 in Xenopus oocytes. P100 exhibits sequence conservation with budding yeast Pat1 and is likely the orthologue of human Pat1a (also called PatL2). P100 is maternally expressed in immature oocytes, but disappears during oocyte maturation. In oocytes, P100 is an RNA binding component of ribosome-free mRNPs, associating with other mRNP components such as Xp54, xRAP55 and CPEB. Translational repression by overexpression of P100 occurred when reporter mRNAs were injected into oocytes. Intriguingly, we found that when P100 was overexpressed in the oocytes, the kinetics of oocyte maturation was considerably retarded. In addition, overexpression of P100 in oocytes significantly affected the accumulation of c-Mos and cyclin B1 during oocyte maturation. These results suggest that P100 plays a role in regulating the translation of specific maternal mRNAs required for the progression of Xenopus oocyte maturation.

DOI： 10.1016/j.ydbio.2010.05.006

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Here we report that structural changes in gonadal basement membranes during sex differentiation in the frog Rana rugosa are revealed using an antibody to its laminin component. Immunohistochemical staining indicated that the first sexual dimorphism appeared in testicular cords and ovarian cavities in differentiating gonads of tadpoles at St. 25-3W, three weeks after they reached St. 25. During development, as the testis enlarged, testicular cord partitions appeared to form by invagination of the testicular epithelium. Ovarian cavities also increased in volume. Laminin-positive basement membranes initially surrounded a partial surface of oocytes close to the ovarian cavity, fully covering growing oocytes by St. X. Laminin-reactive signals were present in somatic cells outside seminiferous tubules in the testis and outside oocytes in one-year-old frogs. BrdU-labeling showed that the number of dividing germ cells increased continuously in male gonads but increased in females only up to St. V, declining at St. X and thereafter. The number of dividing germ cells declined when the basement membranes had fully covered the oocytes. Together, these findings suggest that the first sexual dimorphism in the gonad of R. rugosa first appears as a structural change in the basement membranes. Finally, we speculate that the basement membrane on the surface of oocytes may affect their proliferation in this species.

DOI： 10.1002/jez.607

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Cyp19 is expressed at a high level in the gonad of the female tadpole of the frog Rana rugosa during sex determination. To identify sequence elements important for expression of Cyp19, we isolated a genomic clone (approximately 40 kbp) carrying R. rugosa Cyp19 and analyzed the nucleotide sequence of the 5'-flanking region to search for potential transcription factor binding sites. Sox (SRY-related HMG box) protein and Sf1 binding sites were found in the ovary-specific promoter region of Cyp19. Because Sox3 is located on the sex chromosome in R. rugosa, we conducted the luciferase reporter assay in Xenopus A6 cells using the promoter region. Sox3 drove the reporter gene in the cells, but Sf1 did not. When sequential deletion of the 2.7 kbp Cyp19-promoter region was undertaken, a fragment spanning nucleotides -191 to +48 was sufficient to drive the transcription of the reporter gene. In site-directed mutagenesis, the binding site at -57 in the region was critical for Sox3 responsiveness. Sox3 lacking the HMG box had no ability to promote Cyp19 transcription. In addition, a chromatin immunoprecipitation (ChIP) assay showed that DNA fragments were enriched 8-fold, as determined by real-time PCR, when chromatin was immunoprecipitated with the anti-His antibody against His-tagged Sox3. The results, taken together, suggest that Sox3 activates Cyp19 transcription by its direct binding to the binding site of the Cyp19 promoter region. Sox3 appears to be a factor that directs indifferent gonads to develop into an ovary in R. rugosa.

DOI： 10.1016/j.gene.2009.05.011

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Ascidian is a good model to understand the cellular and molecular mechanisms responsible for mRNA localization with the discovery of a large family of localized maternal mRNAs, called postplasmic/PEM RNAs, which includes more than 40 members in three different ascidian species (Halocynthia roretzi, Ciona intestinalis, and C. savignyi). Among these mRNAs, two types (Type I and Type II) have been identified and show two different localization patterns from fertilization to the eight-cell stage. At the eight-cell stage, both types concentrate to a macromolecular cortical structure called CAB (for Centrosome Attracting Body) in the posterior-vegetal B4.1 blastomeres. The CAB is responsible for unequal cleavages and the partitioning of postplasmic/PEM RNAs at the posterior pole of embryos during cleavage stages. It has also been suggested that the CAB region could contain putative germ granules. In this review, we discuss recent data obtained on the distribution of Type I postplasmic/PEM RNAs from oogenesis to late development, in relation to their localization and translational control. We have first regrouped localization patterns for Type I and Type II into a comparative diagram and included all important definitions in the field. We also have made an exhaustive classification of their embryonic expression profiles (Type I or Type II), and analyzed their functions after knockdown and/or overexpression experiments and the role of the 3'-untranslated region (3'UTR) controlling both their localization and translation. Finally, we propose a speculative model integrating recent data, and we also discuss the relationship between postplasmic/PEM RNAs, posterior specification, and germ cell formation in ascidians.

DOI： 10.1002/dvdy.21109

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Maternal factors, such as a muscle determinant macho-1 mRNA that is localized to the posterior-vegetal cortex (PVC) of fertilized ascidian eggs, are crucial for embryonic axis formation and cell fate specification. Maternal mRNAs that show an identical posterior localization pattern to that of macho-1 in eggs and embryos are called Type I postplasmic/PEM mRNAs. We investigated the functions of five of the nine Type I mRNAs so far known in Halocynthia roretzi: Hr-Wnt-5, Hr-GLUT, Hr-PEM3, Hr-PEN1, and Hr-PEN2. Suppression of their functions with specific antisense morpholino oligonucleotides (MOs) had effects on the formation of various tissues: Hr-Wnt-5 on notochord, muscle, and mesenchyme, although zygotic function of Hr-Wnt-5 is responsible for notochord formation; Hr-GLUT on notochord, mesenchyme, and endoderm; and Hr-PEN2 on muscle, mesenchyme, and endoderm. On the other hand, Hr-PEM3 and Hr-PEN1 MOs seemed to have no effect. We conclude that the functions of at least some localized maternal Type I postplasmic/PEM mRNAs are necessary for early embryonic patterning in ascidians.

DOI： 10.1007/s00427-005-0035-6

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Maternal mRNAs localized to specific regions in eggs play important roles in the establishment of embryonic axes and germ layers in various species. Type I postplasmic/PEM mRNAs, which are localized to the posterior-vegetal cortex (PVC) of fertilized ascidian eggs, such as the muscle determinant macho-1 mRNA, play key roles in embryonic development. In the present study, we analyzed the function of the postplasmic/PEM RNA Hr-POPK-1, which encodes a kinase of Halocynthia roretzi. When the function of POPK-1 was suppressed by morpholino antisense oligonucleotides, the resulting malformed larvae did not form muscle or mesenchyme, as in macho-1-deficient embryos. Epistatic analysis indicated that POPK-1 acts upstream of macho-1. When POPK-1 was knocked down, localization of every Type I postplasmic/PEM mRNA examined, including macho-1, was perturbed, showing diffuse early distribution and eventual concentration into a smaller area. This is the probable reason for the macho-1 dysfunction. The postplasmic/PEM mRNAs such as macho-1 and Hr-PEM1 are co-localized with the cortical endoplasmic reticulum (cER) and move with it after fertilization. Eventually they become highly concentrated into a subcellular structure, the centrosome-attracting body (CAB), at the posterior pole of the cleaving embryos. The suppression of POPK-1 function reduced the size of the domain of concentrated cER at the posterior pole, indicating that POPK-1 is involved in the movement of postplasmic/PEM RNAs via relocalization of cER. The CAB also shrank. These results suggest that Hr-POPK-1 plays roles in concentration and positioning of the cER, as well as in the concentration of CAB materials, such as putative germ plasm, in the posterior blastomeres.

DOI： 10.1242/dev.02049

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The posterior-vegetal cytoplasm (PVC) of fertilized ascidian eggs plays important roles in embryo development. It has been reported that some maternal RNAs are localized to the PVC. We identified four novel type I postplasmic mRNAs that are localized to the PVC through the use of data from a cDNA project of maternal mRNAs in the eggs of Halocynthia roretzi (MAGEST database). The mRNAs are HrGLUT, HrPEN-1, and HrPEM-3, which show similarity to a glucose transporter, a g1-related protein, and Ciona pem-3, respectively; and HrPEN-2, with no similarity. Maternal mRNAs of all four genes were identically localized to the PVC after ooplasmic segregation. During cleavage, they were concentrated in the centrosome-attracting body (CAB) and were then segregated into the small blastomeres located at the posterior pole. This localization pattern is common to all known type I postplasmic mRNAs found so far. HrGLUT, HrPEN-1, and HrPEM-3 were expressed zygotically in various tissues later in embryogenesis: HrGLUT and HrPEM-3 in the mesenchyme and nervous system, and HrPEN-1 in the ectodermal cells.

DOI： 10.1016/S1567-133X(02)00069-8

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