Morpho-anatomical Characterisation and DNA Barcode of Physalis angulata L.
Abstract
Physalis angulata L. is a medicinally important species traditionally used to treat fever, inflammation, and metabolic disorders, making its accurate identification essential for safe pharmaceutical use. This study aims to identify P. angulata through an integrated morpho-anatomical approach and DNA barcoding. A total of 30 individuals from three populations in Central Sulawesi (Parigi Moutong, Sigi, and Donggala) were sampled for morphological characterization, while three representative individuals were used for anatomical analysis and matK sequencing. Morphological traits were assessed for leaves, stems, flowers, and fruits; anatomical features were examined through transverse and paradermal sections using the paraffin method; and DNA barcoding targeted the chloroplast matK region (~700–750 bp). The matK gene was successfully amplified and sequenced for all samples (n = 3), showing 100 % identity and full query coverage with P. angulata accessions in GenBank, and phylogenetic reconstruction clustered all sequences within a single well-supported clade. Morphological data revealed intraspecific variation in leaf and fruit characters, whereas stem and floral traits remained stable across populations. Anatomical observations showed a single-layered epidermis with sinuous cell walls, anomocytic stomata, and collateral vascular bundles with included phloem. Together, these findings provide robust integrative evidence confirming the identity of P. angulata and establish baseline morpho-anatomical and molecular data that can support future pharmacognostic, taxonomic, and conservation research.
References
Ahmed, A. et al., 2023. Gynostemma pentaphyllum an immortal herb with promising therapeutic potential: A comprehensive review on its phytochemistry and pharmacological perspective. International Journal of Food Properties, 26(1), pp.808-832. doi: 10.1080/10942912.2023.2185566.
Ahmed, S. et al., 2022. Pragmatic applications and universality of DNA barcoding for substantial organisms at species level: a review to explore a way forward. BioMed Research International, 2022(1), 1846485. doi: 10.1155/2022/1846485.
Antil, S. et al., 2023. DNA barcoding, an effective tool for species identification: a review. Molecular Biology Reports, 50(1), pp.761-775. doi: 10.1007/s11033-022-08015-7.
Anzani, A.N. et al., 2021. In silico study of DNA barcoding on soka flower (Ixora). Prosiding Biologi Achieving The Sustainable Development Goals With Biodiversity I, 7(1), pp.168-177. doi: 10.24252/psb.v7i1.23693.
Arantes, M.K. et al., 2020. Phenotypic plasticity of leaf anatomical traits helps to explain gas-exchange response to water shortage in grasses of different photosynthetic types. Theoretical and Experimental Plant Physiology, 32(4), pp.341-356.
Azeez, S.O. et al., 2024. Anatomical and molecular characterisation in four Nigerian Physalis (Solanaceae) Species. Nigerian Journal of Botany, 37(2), pp.227-250. doi: 10.4314/njbot.v37i2.6.
Barberon, M., 2017. The endodermis as a checkpoint for nutrients. New phytologist, 213(4), pp.1604-1610. doi: 10.1111/nph.14140.
Bello, A.O. et al., 2017. Leaf epidermal studies of some Solanum (Solanaceae) species in Nigeria. Phytologia Balcanica, 23(1).
Bertolino, L.T. et al., 2019. Impact of stomatal density and morphology on water-use efficiency in a changing world. Frontiers in Plant Science, 10, 225. doi: 10.3389/fpls.2019.00225.
Bhattarai, K. et al., 2025. Improvement of crop production in controlled environment agriculture through breeding. Frontiers in Plant Science, 15, 1524601. doi: 10.3389/fpls.2024.1524601.
Bolboacă, S.D., 2019. Medical diagnostic tests: a review of test anatomy, phases, and statistical treatment of data. Computational and Mathematical Methods in Medicine, 2019(1), 1891569. doi: 10.1155/2019/1891569.
Chorchuhirun, B. et al., 2023. Anatomical Character of Aglaia cucullata (Roxb.) Pellegr., a Medicinal Plant Found in Mangrove Forest of Thailand. Thai Journal of Science and Technology, 11(4), pp.113-128. doi: 10.14456/tjst.2023.10.
Corvalán, L.C. et al., 2025. Is there a key primer for amplification of core land plant DNA barcode regions (rbcL and matK)?. Ecology and Evolution, 15(2), e70961. doi: 10.1002/ece3.70961.
da Silva, H.K. et al., 2024. Selection of morphoagronomic descriptors in Physalis angulata L. using multivariate techniques. Journal of Agricultural Science, 11(1), pp.289-289. doi: 10.5539/jas.v11n1p289.
Deori, M. et al., 2024. A Comprehensive Review on the Impact of Climate Change on Fruit Yield and Quality in Modern Horticultural Practices. International Journal of Plant & Soil Science, 36(1), pp.177-187. doi: 10.9734/ijpss/2024/v36i14348.
Dini, S. et al., 2022. Phytochemical and biological activities of some Iranian medicinal plants. Pharmaceutical Biology, 60(1), pp.664-689. doi: 10.1080/13880209.2022.2046112.
Do, H.D.K. et al., 2019. The newly developed single nucleotide polymorphism (SNP) markers forapotent medicinal plant, Crepidiastrum denticulatum (Asteraceae), were inferred fromcomplete chloroplast genome data. Molecular Biology Reports, 46(3), pp.3287-3297. doi: 10.1007/s11033-019-04789-5.
Driesen, E. et al., 2020. Influence of environmental factors light, CO2, temperature, and relative humidity on stomatal opening and development: A review. Agronomy, 10(12), 1975. doi: 10.3390/agronomy10121975.
Ebigwai, J.K. et al., 2020. Application of Maturase K Primer to Determining Nomenclatural identities of Four Morphological Similar Mimosoid Clade Members. Life Science Journal, 17(6). doi: 10.7537/marslsj170620.07.
Elhawary, E.A. et al., 2025. Seasonal variation effect on different Physalis peruviana L. (Solanaceae) waste extracts and investigation of their efficacy against Culex pipiens and Musca domestica. Scientific Reports, 15(1), 20231. doi: 10.1038/s41598-025-89854-9.
Fan, Y. et al., 2007. Patterns of insertion and deletion in mammalian genomes. Curr Genomics, 8(6), pp.370-378. doi: 10.2174/138920207783406479.
Fathiya, N. et al., 2018. Molecular identification of Shorea johorensis in Ketambe Research Station, Gunung Leuser National Park. Jurnal Natural, 18(2), pp.56-64. doi: 10.24815/jn.v18i2.10123.
Firdaus, A. et al., 2022. Phenological of cutleaf groundcherry (Physalis angulata L.) based on BBCH scale. Jurnal Agronomi Tanaman Tropika, 4(2), pp.241-254. doi: 10.36378/juatika.v4i2.2063.
Flint-Garcia, S.A., 2013. Genetics and consequences of crop domestication. Journal of Agricultural and Food Chemistry, 61(35), pp.8267-8276. doi: 10.1021/jf305511d.
García-Verdugo, C. et al., 2019. How repeatable is microevolution on islands? Patterns of dispersal and colonization-related plant traits in a phylogeographical context. Annals of Botany, 123(3), pp.557-568. doi: 10.1093/aob/mcy191.
Garibyan, L. & Avashia, N., 2013. Polymerase chain reaction. Journal of Investigative Dermatology, 133(3), pp.1-4. doi: 10.1038/jid.2013.1.
Gray, A. et al., 2020. Flanking support: how subsidiary cells contribute to stomatal form and function. Frontiers in Plant Science, 11, 881. doi: 10.3389/fpls.2020.00881.
Harrison, E.L. et al., 2020. The influence of stomatal morphology and distribution on photosynthetic gas exchange. The Plant Journal, 101(4), pp.768-779. doi: 10.1111/tpj.14560.
Hasanuddin & Fitriana., 2014. Relationship fenetik 12 species of Asteraceae family members. Journal EduBio Tropical, 2(2), pp.187-250.
Ho, V.T. et al., 2021. Comparison of matK and rbcL DNA barcodes for genetic classification of jewel orchid accessions in Vietnam. Journal of Genetic Engineering and Biotechnology, 19(1), 93. doi: 10.1186/s43141-021-00188-1.
Hong, Y. et al., 2021. ENJ algorithm can construct triple phylogenetic trees. Molecular Therapy Nucleic Acids, 23, pp.286-293. doi: 10.1016/j.omtn.2020.11.004.
Ilham, M. et al., 2022. Morpho-anatomical characterizationand DNA barcoding of Achillea millefolium L. Biodiversitas, 23(4), pp.1958-1969. doi: 10.13057/biodiv/d230430.
Kayama, K. et al., 2021. Prediction of PCR amplification from primer and template sequences using recurrent neural network. Scientific Reports, 11, 7493. doi: 10.1038/s41598-021-86357-1.
Keskin, E. & Atar, H.H., 2013. DNA barcoding commercially important fish species of Turkey. Molecular Ecology Resources, 13(5), pp.788-797. doi: 10.1111/1755-0998.12120.
Khan, R. et al., 2017. Scanning electron and light microscopy of foliar epidermal characters: A tool for plant taxonomists in the identification of grasses. Microscopy Research and Technique, 80(10), pp.1123-1140. doi: 10.1002/jemt.22909.
Khan, S.U. et al., 2013. Morpho-anatomical study of selected plants of district bannu, khyber pakhtunkhwa, pakistan. Pakistan Journal of Weed Science Research, 19(4).
Knapp, S. et al., 2013. Wild relatives of the eggplant (Solanum melongena L.: Solanaceae): new understanding of species names in a complex group. PLoS One, 8(2), e57039. doi: 10.1371/journal.pone.0057039.
Kumar, P. et al., 2022. Comparative morpho-anatomical standardization and chemical profiling of root drugs for distinction of fourteen species of family Apocynaceae. Botanical Studies, 63(1), 12. doi: 10.1186/s40529-022-00342-z.
Kumar, S. et al., 2018. MEGA X: Molecular evolutionary genetics analysis across computing platforms. Molecular Biology and Evolution, 35(6), pp.1547-1549. doi: 10.1093/molbev/msy096.
Kurniawati, S. & Hartati, N.S., 2018. Optimization of the annealing temperature with degenerate primer for amplifiation of arginine decarboxylase (ADC) fragment gene from genomic DNA of Maluku Tenggara local cassava. Jurnal Ilmu Dasar, 19(2), pp.135-142. doi: 10.19184/jid.v19i2.6261.
Leelavathy, S. & Sankar, P.D., 2021. Maturase K (matK) as a Barcode in Bamboos. Research Journal of Pharmacy and Technology, 14(2), pp.955-958. doi: 10.5958/0974-360X.2021.00170.0.
Lemoine, F. et al., 2018. Renewing Felsenstein’s phylogenetic bootstrap in the era of big data. Nature, 556(7702), pp.452-456.
Lonard, R.I. et al., 2017. The biological flora of coastal dunes and wetlands: Avicennia germinans (L.) L. Journal of Coastal Research, 33(1), pp.191-207. doi: 10.2112/JCOASTRES-D-16-00013.1.
Madhavan, C. et al., 2025. Anatomical adaptations of mangroves to the intertidal environment and their dynamic responses to various stresses. Biological Reviews, 100(3), pp.1019-1046. doi: 10.1111/brv.13172.
Makhmudova, M.M. et al., 2022. Anatomical structure of vegetative organs of medicinal plants Physalis angulata L. and their chemical composition. The American Journal of Agriculture and Biomedical Engineering, 4(01), pp.13-21. doi: 10.37547/tajabe/Volume04Issue01-03.
Malik, O.A. et al., 2022. Automated real-time identification of medicinal plants species in natural environment using deep learning models a case study from Borneo Region. Plants, 11(15), 1952. doi: 10.3390/plants11151952.
Maruzy, A. et al., 2020. Comparatison study of macroscopic and microscopic characters in three species Phyllanthus L. Floribunda, 6(4), pp.154-166. doi: 10.32556/floribunda.v6i4.2020.312.
Medda, S. et al., 2022. Influence of climate change on metabolism and biological characteristics in perennial woody fruit crops in the Mediterranean environment. Horticulturae, 8(4), 273. doi: 10.3390/horticulturae8040273.
Michel, C.I. et al., 2016. The nuclear internal transcribed spacer (ITS2) as a practical plant DNA barcode for herbal medicines. Journal of Applied Research on Medicinal and Aromatic Plants, 3(3), pp.94-100. doi: 10.1016/j.jarmap.2016.02.002.
Mirzaee, F. et al., 2019. Therapeutic activities and phytochemistry of Physalis species based on traditional and modern medicine. Research Journal of Pharmacognosy, 6(4), pp.79-96. doi: 10.22127/rjp.2019.93529.
Ni, X. et al., 2022. Variation and determinants of leaf anatomical traits from boreal to tropical forests in eastern China. Ecological Indicators, 140, 108992. doi: 10.1016/j.ecolind.2022.108992.
Novitasari, A. et al., 2024. Physalis angulata Linn. as a medicinal plant. Biomedical Reports, 20(3), pp.1-16. doi: 10.3892/br.2024.1735.
Oliva-Ruiz, M. et al., 2023. Growth, phenotypic plasticity and fruit quality in tomato: a study under high temperature and elevated CO2. Horticulturae, 9(12), 1266. doi: 10.1016/j.heliyon.2021.e05988.
Prasetyawan, F. et al., 2024. Morphological analysis of ciplukan plant (Physalis angulata L.) pollen for macroscopic identification. International Journal of Science and Environment (IJSE), 4(1), pp.11-17.
Purnobasuki, H. et al., 2017. Morphology of four root types and anatomy of root-root junction in relation gas pathway of Avicennia marina (Forsk) Vierh roots. Vegetos, 30(2), pp.100-104. doi: 10.5958/2229-4473.2017.00143.4.
Ragab, O.G. et al., 2022. Micromorphological characteristics of leaf epidermis for the identification of certain Solanoideae (Solanaceae). International Journal of Theoretical and Applied Research, 1(1), pp.38-48. doi: 10.21608/ijtar.2022.139481.1002
Rajput, K.S. et al., 2022. Inter-and intraxylary phloem in vascular plants: A review of subtypes, occurrences, and development. Forests, 13(12), 2174. doi: 10.3390/f13122174.
Rashid, P. et al., 2020. Leaf anatomical adaptation of eighteen mangrove plant species from the Sundarbans in Bangladesh. Bangladesh Journal of Botany, 49(4), pp.903-911. doi: 10.3329/bjb.v49i4.52496.
Raymundo, C.E. et al., 2025. Heteroblasty in Conchocarpus heterophyllus (A. St.-Hil.) Kallunki & Pirani (Rutaceae): An approach of leaf development from the unifoliolate leaves. Flora, 322, 152645. doi: 10.1016/j.flora.2024.152645.
Rivera, D.E. et al., 2019. A screening of plants used in Colombian traditional medicine revealed the anti-inflammatory potential of Physalis angulata calyces. Saudi Journal of Biological Sciences, 26(7), pp.1758-1766. doi: 10.1016/j.sjbs.2018.05.030.
Rodrigues, M.H.B.S. et al., 2021. Characterization of physiological maturity of Physalis peruviana L. fruits. Semina: Ciências Agrárias Londrina, 42(3), pp.929-948. doi: 10.5433/1679-0359.2021v42n3p929
Rouhan, G. & Gaudeul, M., 2020. Plant taxonomy: A historical perspective, current challenges, and perspectives. Molecular Plant Taxonomy: Methods and Protocols, pp.1-38. doi: 10.1007/978-1-0716-0997-2_1.
Santos, K.S.D. et al., 2022. Genetic variability of Physalis ixocarpa and P. philadelphica from physicochemical fruit traits. Pesquisa Agropecuária Brasileira, 56, e01534. doi: 10.1590/s1678-3921.pab2021.v56.01534.
Sanyal, G. et al., 2015. Insertion-deletion as informative characters in DNA barcoding. International Journal of Multimedia and Ubiquitous Engineering, 10(10), pp.67–74. doi: 10.14257/ijmue.2015.10.10.07.
Schneider, R.F. & Meyer, A., 2017. How plasticity, genetic assimilation and cryptic genetic variation may contribute to adaptive radiations. Molecular ecology, 26(1), pp.330-350. doi: 10.1111/mec.13880.
Seleem, E.A. & Nassar, R., 2021. Morphological and anatomical studies on Physalis peruviana L. and Physalis ixocarpa Brot. Exhornem. Journal of Plant Production, 12(11), pp.1179-1183. doi: 10.21608/jpp.2021.100328.1069.
Silva, J.C. et al., 2024. Comparative leaf anatomy of two species of Ipomoea L. (Convolvulaceae): taxonomic importance and adaptations to xeric conditions of the cangas. PeerJ, 12, e18599. doi: 10.7717/peerj.18599.
Singirala, S.K. et al., 2025. Extraction of bioactive compounds from Withania somnifera: The biological activities and potential application in the food industry: A Review. International Journal of Food Science, 2025(1), 9922626. doi: 10.1155/ijfo/9922626.
Smith, D.R. et al., 2011. The GC-rich mitochondrial and plastid genomes of the green alga Coccomyxa give insight into the evolution of organelle DNA nucleotide landscape. PLoS One, 6(8), e23624. doi: 10.1371/journal.pone.0023624.
Stefi, A.L. et al., 2025. Oregano young plants cultured at low temperature reveal an enhanced healing effect of their extracts: anatomical, physiological and cytotoxicity approach. Metabolites, 15(2), 103. doi: 10.3390/metabo15020103.
Susetyarini, E. et al., 2020. The identification of morphological and anatomical structures of Pluchea indica. Journal of Physics: Conference Series (JPCS), 1539, 012001. doi: 10.1088/1742-6596/1539/1/012001.
Suwanphakdee, C. et al., 2024. Characterization of anatomical characters of Peperomia (Piperaceae) from Asia for taxonomy. Botanical Journal of the Linnean Society, 205(3), pp.268-291. doi: 10.1093/botlinnean/boad075.
Tindi, M. et al., 2017. The DNA barcode and molecular phylogenetic analysis several bivalve species from North Sulawesi waters based on COI gene. Jurnal Pesisir dan Laut Tropis, 1(2), pp.33-38. doi: 10.35800/jplt.5.2.2017.15050.
Wahyuni, D.K. et al., 2019. Morpho-anatomical structure and DNA barcode of Sonchus arvensis L. Biodiversitas Journal of Biological Diversity, 20(8), pp.2417-2426. doi: 10.13057/biodiv/d200841.
Wahyuni, D.K. et al., 2022. Morpho-anatomical characterization and DNA barcoding analysis of Pluchea indica (L.) Less. Biodiversitas Journal of Biological Diversity, 23(8). doi: 0.13057/biodiv/d230851.
Wahyuni, D.K. et al., 2024. Morpho-anatomical characterization and DNA barcoding of Artemesia vulgaris L. Brazilian Journal of Biology, 84, e278393. doi: 10.1590/1519-6984.278393.
Wang, M. et al., 2024. From species to varieties: How modern sequencing technologies are shaping Medicinal Plant Identification. Genes, 16(1), 16. doi: 10.3390/genes16010016.
Yang, J. et al., 2017. Development of chloroplast and nuclear DNA markers for Chinese oaks (Quercus Subgenus Quercus) and assessment of their utility as DNA Barcodes. Frontiers in Plant Science, 8, 816. doi: 10.3389/fpls.2017.00816.
Yue, J. et al., 2021. Application of identification and evaluation techniques for ethnobotanical medicinal plant of genus Panax: A review. Critical Reviews in Analytical Chemistry, 51(4), pp.373-398. doi: 10.1080/10408347.2020.1736506.
Zhang, X. et al., 2021. Quantitative extraction and analysis of pear fruit spot phenotypes based on image recognition. Computers and Electronics in Agriculture, 190, 106474. doi: 10.1016/j.compag.2021.106474.
