{"title":"Typical tetra-mediated signaling and plant architectural changes regulate salt-stress tolerance in indica rice genotypes.","authors":"Shivani Shivani, Rohit Ghosh, Adinpunya Mitra, Arpita Das, Joydeep Banerjee","doi":"10.1007/s00709-025-02035-3","DOIUrl":"10.1007/s00709-025-02035-3","url":null,"abstract":"<p><p>Upon exposure to salt stress, calcium signaling in plants activates various stress-responsive genes and proteins along with enhancement in antioxidant defense to eventually regulate the cellular homeostasis for reducing cytosolic sodium levels. The coordination among the calcium signaling molecules and transporters plays a crucial role in salinity tolerance. In the present study, twenty-one diverse indigenous rice genotypes were evaluated for salt tolerance during the early seedling stage, and out of that nine genotypes were further selected for physio-biochemical study. Further analysis identified potential salt-tolerant and salt-sensitive genotypes with tolerant lines exhibiting lower Na<sup>+</sup>/K<sup>+</sup> ratio. Plant phenotype clearly documented the root architectural changes among the most salt-tolerant and sensitive genotypes, while the histo-biochemical DAB and NBT staining and in-gel SOD activity clearly revealed the differential ROS accumulation between the contrasting genotypes and their antioxidant activity. Ultrastructural study depicted stronger UV autofluorescence in the hypodermal and vascular bundle regions, while phloroglucinol staining displayed intense coloration in the vascular bundle region of Bhutmuri, the salt-tolerant genotype, compared to Manipuri Black, the salt-sensitive one showing differential lignification among the contrasting genotype to combat salt stress. Based on expression study, our proposed model depicted immediate upregulation (short-term) of OsSOS3 and OsNHX1 along with gradual upregulation of OsHKT and downregulation of OsSOS1 throughout the stress period to protect the tolerant plant through signaling cascade, while the inadequate upregulation of OsSOS3, OsNHX1, and OsHKT under early stress, coupled with poor coordination between the expression of OsSOS1 and OsSOS3 genes, makes the plant salt sensitive.</p>","PeriodicalId":20731,"journal":{"name":"Protoplasma","volume":" ","pages":"837-855"},"PeriodicalIF":2.5,"publicationDate":"2025-07-01","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143053425","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
ProtoplasmaPub Date : 2025-07-01Epub Date: 2025-02-03DOI: 10.1007/s00709-025-02039-z
Kadri Oumaima, Mohammad Shakhawat Hossain, Wenxiu Ye, Eiji Okuma, Mohammad Issak, Mohammad Mahbub Islam, Misugi Uraji, Yoshimasa Nakamura, Izumi C Mori, Shintaro Munemasa, Yoshiyuki Murata
{"title":"TGG1 and TGG2 mutations impair allyl isothiocyanate-mediated stomatal closure in Arabidopsis thaliana.","authors":"Kadri Oumaima, Mohammad Shakhawat Hossain, Wenxiu Ye, Eiji Okuma, Mohammad Issak, Mohammad Mahbub Islam, Misugi Uraji, Yoshimasa Nakamura, Izumi C Mori, Shintaro Munemasa, Yoshiyuki Murata","doi":"10.1007/s00709-025-02039-z","DOIUrl":"10.1007/s00709-025-02039-z","url":null,"abstract":"<p><p>Myrosinase, referred to as thioglucoside glucohydrolase (TGG), plays a crucial role in plant physiology through catalyzing the hydrolysis of glucosinolates into bioactive isothiocyanates. In Arabidopsis thaliana, the myrosinases TGG1 and TGG2 are essential for abscisic acid- and methyl jasmonate-induced stomata closure. Allyl isothiocyanate (AITC), one of myrosinase products, triggers stomatal closure in A. thaliana. We investigated stomatal responses to AITC to clarify the role of TGG1 and TGG2 in Arabidopsis guard-cell signaling. Allyl isothiocyanate at 50 μM and 100 μM induced stomatal closure in the tgg1 and tgg2 single mutants but not in the tgg1 tgg2 double mutant. Furthermore, AITC at 50 μM induced the production of reactive oxygen species and nitric oxide, cytosolic alkalization, and oscillations in cytosolic free calcium concentration in guard cells of both wild-type and mutant plants. These findings suggest that TGG1 and TGG2 are involved in AITC signaling pathway through interaction with signal component(s) downstream of these signaling events, which is not accompanied by hydrolysis of glucosinolates because of the difference in subcellular localization between enzymes (myrosinases) and substrates (glucosinolates).</p>","PeriodicalId":20731,"journal":{"name":"Protoplasma","volume":" ","pages":"1023-1027"},"PeriodicalIF":2.5,"publicationDate":"2025-07-01","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://www.ncbi.nlm.nih.gov/pmc/articles/PMC12141395/pdf/","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143080751","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"OA","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
ProtoplasmaPub Date : 2025-07-01Epub Date: 2025-02-18DOI: 10.1007/s00709-025-02041-5
Santiago Michavila, Antonio Encina, Alfonso G De la Rubia, María Luz Centeno, Penélope García-Angulo
{"title":"An immunohistochemical approach to cell wall polysaccharide specialization in maritime pine (Pinus pinaster) needles.","authors":"Santiago Michavila, Antonio Encina, Alfonso G De la Rubia, María Luz Centeno, Penélope García-Angulo","doi":"10.1007/s00709-025-02041-5","DOIUrl":"10.1007/s00709-025-02041-5","url":null,"abstract":"<p><p>Pine needles are specialized leaves bearing distinctive anatomical features whose function is to minimize water loss. At the molecular level, needle specialization is expected to lead to heterogeneity in cell wall (CW) composition. By immunohistochemical analysis of CW epitopes in maritime pine (Pinus pinaster) needles, we described the variability of CW composition in all tissues, discussing its possible relationship with the cell function. Cells specialized in water transport, tracheids, transfusion tracheids, and endodermis, together with epi/hypodermal and stomatal cells, had similar CW composition, where epitopes for pectins (homogalacturonan and rhamnogalacturonan-I), xyloglucan, and heteroxylans were abundant. Heteroxylans differ between the dermis, enriched in arabinoxylan, and endodermis, enriched in glucuronoxylan. Xylem CWs were very rich in (arabino)xylans and homogalacturonan and distinctively lacked immunolabelling for rhamnogalacturonan-I. CWs from phloem, transfusion phloem, mesophyll, and resin ducts were enriched in homogalacturonan, rhamnogalacturonan-I, and xyloglucan, showing a low content for heteroxylan epitopes. Arabinogalactan proteins were detected in mesophyll and in cells specialized in solute transport such as phloem, duct sheath, and guard cells. These results show that tissues with similar functions have similar polysaccharide composition, shedding light on the role of CW components through different tissues.</p>","PeriodicalId":20731,"journal":{"name":"Protoplasma","volume":" ","pages":"979-991"},"PeriodicalIF":2.5,"publicationDate":"2025-07-01","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://www.ncbi.nlm.nih.gov/pmc/articles/PMC12141410/pdf/","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143441803","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"OA","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"Genome-wide association mapping for stay-green and stem reserve mobilization traits in wheat (Triticum aestivum L.) under combined heat and drought stress.","authors":"Animireddy China Malakondaiah, Ajay Arora, Hari Krishna, Sukumar Taria, Sudhir Kumar, Narayana Bhat Devate, Jasdeep Chatrath Padaria, Sekar Kousalya, Sahana Police Patil, Pradeep Kumar Singh","doi":"10.1007/s00709-025-02031-7","DOIUrl":"10.1007/s00709-025-02031-7","url":null,"abstract":"<p><p>Stay-green (SG) and stem reserve mobilization (SRM) are two significant mutually exclusive traits, which contributes to grain-filling during drought and heat stress in wheat. The current research was conducted in a genome-wide association study (GWAS) panel consisting of 278 wheat genotypes of advanced breeding lines to find the markers linked with SG and SRM traits and also to screen the superior genotypes. SG and SRM traits, viz. soil plant analysis development (SPAD) value, canopy temperature (CT), normalized difference vegetation index (NDVI), leaf senescence rate (LSR) and stem reserve mobilization efficiency (SRE) were recorded. The trial was conducted in α-lattice design, under control and combined heat and drought stress (HD). Analysis of variance and descriptive statistics showed a significant difference across the evaluated traits. The highest mean of SRE (31.7%) and SRM (0.42 g/stem) was reported in HD, while highest SRE in HD and lowest in control was 52.56% and 15.7%, respectively. Genotyping was carried out using the 35 K Axiom R Wheat Breeder's Array, 14,625 SNPs were kept after filtering. Through GWAS, 36 significant marker trait associations (MTAs) were identified on 16 distinct chromosomes; out of this, 22 MTAs were found under control and 14 MTAs under HD. Candidate genes that code for UDP-glycosyltransferase 73C4-like and protein detoxification 40-like was linked to SPAD and CT respectively. One MTAs was detected for SRM on chromosome 6B that code for wall associated receptor kinase 4 like. These SNPs can be utilized to generate cultivars that adapt to climate change by a marker-assisted gene transfer.</p>","PeriodicalId":20731,"journal":{"name":"Protoplasma","volume":" ","pages":"759-778"},"PeriodicalIF":2.5,"publicationDate":"2025-07-01","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"142979791","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"Successful application of microspore culture by adding kinetin and methylene blue to improve microspore embryogenesis in Chinese kale breeding.","authors":"Haonan Li, Zhe Wang, Weina Zhou, Jinyan Li, Zifan Zhao, Chao Zheng, Zhiyong Liu, Yun Zhang","doi":"10.1007/s00709-025-02032-6","DOIUrl":"10.1007/s00709-025-02032-6","url":null,"abstract":"<p><p>Microspore culture is an efficient and rapid method that produces doubled haploid (DH) lines for hybrid breeding in crops and vegetables. However, the low frequency of microspore embryogenesis and spontaneous diploidization in Chinese kale still require improvement. In the present work, an efficient microspore culture protocol was constructed and used for DH producing in Chinese kale breeding. The results showed that kinetin (KT) and methylene blue (MB) improved microspore embryogenesis and embryo development. The spontaneous diploidization rates were below 14.48%. Therefore, artificial doubling was used for haploids; the doubled haploid rate was significantly increased, and the highest doubling rate reached 33.75% with pendimethalin. 98 DH lines were obtained in this study. This study also identified the horticultural characters of the DH lines, which showed significant differences between them. All DH lines were self-incompatible. The hybrid crosses were performed using five DH lines selected for the Chinese kale breeding. This study was helpful to accelerate the application of microspore culture and provide DH lines for Chinese kale hybrid breeding.</p>","PeriodicalId":20731,"journal":{"name":"Protoplasma","volume":" ","pages":"801-813"},"PeriodicalIF":2.5,"publicationDate":"2025-07-01","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143010567","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
ProtoplasmaPub Date : 2025-07-01Epub Date: 2025-01-30DOI: 10.1007/s00709-025-02036-2
Jéssica Nascimento, Mariela Sader, Tiago Ribeiro, Andrea Pedrosa-Harand
{"title":"Influence of Ty3/gypsy and Ty1/copia LTR-retrotransposons on the large genomes of Alstroemeriaceae: genome landscape of Bomarea edulis (Tussac) Herb.","authors":"Jéssica Nascimento, Mariela Sader, Tiago Ribeiro, Andrea Pedrosa-Harand","doi":"10.1007/s00709-025-02036-2","DOIUrl":"10.1007/s00709-025-02036-2","url":null,"abstract":"<p><p>Repetitive elements are the main components of many plant genomes and play a crucial role in the variation of genome size and structure, ultimately impacting species diversification and adaptation. Alstroemeriaceae exhibits species with large genomes, not attributed to polyploidy. In this study, we analysed the repetitive fraction of the genome of Bomarea edulis through low-coverage sequencing and in silico characterization, and compared it to the repeats of Alstroemeria longistaminea, a species from a sister genus that has been previously characterized. LTR-retrotransposons were identified as the most abundant elements in the B. edulis genome (50.22%), with significant variations in abundance for specific lineages between the two species. The expansion of the B. edulis genome was likely due to three main lineages of LTR retrotransposons, Ty3/gypsy Tekay and Retand and Ty1/copia SIRE, all represented by truncated elements which were probably active in the past. Furthermore, the proportion of satDNA (~ 7%) was six times higher in B. edulis compared to A. longistaminea, with most families exhibiting a dispersed, uniform distribution in the genome. SatDNAs, thus, contributed to some extent to genome obesity. Despite diverging around 29 Mya, both species still share some satDNA families and retrotransposons. However, differences in repeat abundances and sequence variants led to genome differentiation despite their similar sizes and structure.</p>","PeriodicalId":20731,"journal":{"name":"Protoplasma","volume":" ","pages":"881-894"},"PeriodicalIF":2.5,"publicationDate":"2025-07-01","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143067523","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
ProtoplasmaPub Date : 2025-07-01Epub Date: 2025-02-16DOI: 10.1007/s00709-025-02034-4
Juan Liu, Yuangang Zhu, Xinyue Liu, Jian Song, Ligang Tang, Liang Shen, Zhongmin Dai
{"title":"Morphological development of the endosperm epidermal cells in waxy wheat cultivars.","authors":"Juan Liu, Yuangang Zhu, Xinyue Liu, Jian Song, Ligang Tang, Liang Shen, Zhongmin Dai","doi":"10.1007/s00709-025-02034-4","DOIUrl":"10.1007/s00709-025-02034-4","url":null,"abstract":"<p><p>Endosperm epidermal cells (EECs) accumulate large quantities of nutrients; they also play key roles in facilitating solute transport. Comprehensive knowledge about the dynamic development of EECs is needed to understand the relationship between their dual functions. In this study, the developmental characteristics of EECs in wheat grains of two near-isogenic lines (Shimai19-P and Shimai19-N) and in the parent wheat cultivar Shimai19 were compared using light and scanning electron microscopy. The intermediate EECs located adjacent to the nucellar projection (NP) on the ventral surface of wheat grains rapidly differentiated. Eight days after pollination (8 DAP), these EECs were larger in Shimai19-N than in the other wheat cultivars; they had differentiated into endosperm transfer cells (ETCs). At 14 DAP, the number of ETCs reached a maximum and then gradually decreased in all three wheat varieties. The lateral ETCs and the ETCs on both sides of the crease were longer than ACs; they reached their maximum length at 16 DAP, becoming gradually shorter thereafter. The dorsal ACs became increasingly thicker during wheat grain development. Overall, these results suggested that EECs near the EC and crease are important for efficient nutrient transport, whereas EECs in other regions of wheat grains mainly play a role in nutrient storage.</p>","PeriodicalId":20731,"journal":{"name":"Protoplasma","volume":" ","pages":"957-977"},"PeriodicalIF":2.5,"publicationDate":"2025-07-01","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143426031","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
ProtoplasmaPub Date : 2025-07-01Epub Date: 2025-02-19DOI: 10.1007/s00709-025-02045-1
O V Nakonechnaya, Yu A Khrolenko, A V Kalachev, V M Kazarin, T Y Gorpenchenko
{"title":"Structural characteristics of leaf trichomes and epidermis in species from three sections of the genus Aristolochia.","authors":"O V Nakonechnaya, Yu A Khrolenko, A V Kalachev, V M Kazarin, T Y Gorpenchenko","doi":"10.1007/s00709-025-02045-1","DOIUrl":"10.1007/s00709-025-02045-1","url":null,"abstract":"<p><p>The structure of leaf epidermis in species from three sections (Diplolobus, Gymnolobus, and Siphisia) of the genus Aristolochia has been examined by light and scanning microscopy methods. As the results show, the number of the stomata and the length of stomatal guard cells in plants growing ex situ are not related to habitat conditions but are probably determined genetically. According to the type of leaf epicuticular waxes, the studied taxa can be divided into three groups, of which the Aristolochia-type is the most common. This type is characteristic of all studied species of the section Siphisia. The presence of two types of trichomes in members of this section distinguishes them from the other sections. Representatives of the sections with the Convallaria-type wax lack trichomes. The noted differences allow accurate identification of each of the species in the genus. The features of the epidermis surface, types and arrangement of stomata, and types of trichomes are additional diagnostic characteristics useful for taxonomy.</p>","PeriodicalId":20731,"journal":{"name":"Protoplasma","volume":" ","pages":"993-1007"},"PeriodicalIF":2.5,"publicationDate":"2025-07-01","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143449958","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"Elucidating the wedelolactone biosynthesis pathway from Eclipta prostrata (L.) L.: a comprehensive analysis integrating de novo comparative transcriptomics, metabolomics, and molecular docking of targeted proteins.","authors":"Anjum Bano, Archana Kumari, Akansha Pandey, Akhilesh Kumar, Mallika Madan, Anshu Mohanta, Emma Anjali Minj, Tushar Pandey, Sanjeev Kanojiya, Richa Pandey, Rakesh Kumar Shukla, Vineeta Tripathi","doi":"10.1007/s00709-025-02030-8","DOIUrl":"10.1007/s00709-025-02030-8","url":null,"abstract":"<p><p>Eclipta prostrata belongs to the Asteraceae family. The plant contains bioactive compounds like wedelolactone (WDL) and demethylwedelolactone (DW). Its transcriptomic information engaged with secondary metabolite biosynthesis is not available. Based on differential accumulation of WDL and DW in root, shoot of the mature plant, we performed comparative de novo transcriptome of root and shoot tissue in three independent biological replicates and generated 49820 unique transcripts. Annotation resulted in significant matches for 43,015 unigenes. Based on differential gene expression data, we found WDL biosynthesis-related transcripts, which were mainly upregulated in shoot. Finally, 13 selected differentially expressed transcripts related to WDL biosynthesis that were validated by qRT-PCR. Detailed tissue-specific metabolite and transcript profiling revealed that DW highly accumulated in root and WDL accumulation was high in aerial part along with transcripts. For WDL pathway exploration, we did integrated profiling of 08 metabolites and 13 transcripts and witnessed that only naringenin, apigenin, DW, and WDL were detected in different developmental stages. Taking leads from the findings, we postulated that naringenin to apigenin pathway is one potential route for WDL biosynthesis. Moreover, wound stress led to accumulation of DW and WDL and related biosynthetic transcripts. Furthermore, the selected enzymes were subjected to molecular docking and binding studies for the predicted substrates involved in crucial and advance steps of WDL biosynthesis. A comprehensive analysis integrating de novo transcriptomics, metabolomics, and molecular docking of targeted proteins paves the way for the elucidation of the putative wedelolactone biosynthesis pathway from E. prostrata.</p>","PeriodicalId":20731,"journal":{"name":"Protoplasma","volume":" ","pages":"815-835"},"PeriodicalIF":2.5,"publicationDate":"2025-07-01","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143024384","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
ProtoplasmaPub Date : 2025-07-01Epub Date: 2025-02-10DOI: 10.1007/s00709-025-02040-6
A Bonfitto, R Randi, M Ciubotaru, L Alibardi
{"title":"Microscopic and ultrastructural observations on the regenerating scales of the lizard Podarcis muralis clarify the origin of the micro-ornamentation.","authors":"A Bonfitto, R Randi, M Ciubotaru, L Alibardi","doi":"10.1007/s00709-025-02040-6","DOIUrl":"10.1007/s00709-025-02040-6","url":null,"abstract":"<p><p>The present study conducted using immunofluorescence, scanning electron microscopy, and transmission electron microscopy aims to determine the origin of the lamellated pattern and of the micro-ornamentation formed in tail scales of the lacertid lizard Podarcis muralis. Regenerating scales shows the typical formation of all the epidermal layers of lizard epidermis, including clear and Oberhautchen layers. The latter initiates the accumulation of hard corneous material containing corneous beta proteins (CBPs) that determine a tension with the overlaying softer clear layer containing mainly intermediate filaments of keratins (IFKs). The two layers, initially joined by numerous desmosomes, are later displaced one from the other with the growth of regenerating scales, forming a slanted surface as observed under transmission electron microscopy (TEM). At the beginning of scale regeneration, the slanted tips form an irregular lamellated pattern on the surface of Oberhautchen cells, appearing as crests or waves under scanning electron microscopy (SEM). In the following scale differentiation, growth, and shedding of molt, the irregular crests form a more ordered and parallel microsculptured and micro-ornamentation pattern when Oberhautchen and beta-cells merge one to another and give rise to a mature beta-layer. Hard CBP-based corneous material and electron-dense materials of unknown composition together with IFKs are accumulated in the slanted surface of the differentiating Oberhautchen cells. During scale growth, the Oberhautchen surface matures into a jig-saw outline that gives rise to the lamellated pattern of mature micro-ornamentation. The study suggests that complex micro-ornamentation patterns in other lizard species can also vary during scale formation, in development, growth, or regeneration.</p>","PeriodicalId":20731,"journal":{"name":"Protoplasma","volume":" ","pages":"935-955"},"PeriodicalIF":2.5,"publicationDate":"2025-07-01","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143382516","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}