Anatomical traits and economic potential of Amorphophallus variabilis and conservation implications in Selo Arjuno Forest, Central Java, Indonesia
Main Article Content
Abstract
Abstract. Wahidah BF, Khasanah R, Husain F. 2026. Anatomical traits and economic potential of Amorphophallus variabilis and conservation implications in Selo Arjuno Forest, Central Java, Indonesia. Biodiversitas 27 (2): d270213. https://doi.org/10.13057/biodiv/d270213. Amorphophallus variabilis is rarely utilized or studied despite its ecological and economic value. This study aimed to identify its anatomical characteristics, secretory structure, and the distribution of ergastic substances in its organs, including assessing their implications for conservation and local economic development. The study was conducted in Selo Arjuno Forest, Kendal District, Central Java, Indonesia. Anatomical slides from six mature individuals were prepared with paraffin methods and examined under a light microscope. Socio-economic data were collected through in-depth interviews. All data were analysed descriptively. The results showed that tubers are the primary storage organs, with a very high starch granule content (±8400/mm²), far exceeding those in petioles (±25/mm²) and leaf blades (±71/mm²). Sand and druse calcium oxalate crystals dominate tubers, contributing to stress tolerance. Tannin distribution is organ-specific, namely schizogenous cavities are more abundant in petioles, while idioblasts occur more frequently in tubers. Interviews indicate that A. variabilis is rarely cultivated, although its glucomannan-rich tubers have strong potential for functional food and health products. These findings highlight the importance of functional anatomy for conservation planning. In situ conservation and community-based ex situ cultivation can support sustainable management while creating utilization opportunities that enhance local livelihoods.
Article Details
Issue
Section

This work is licensed under a Creative Commons Attribution-NonCommercial-ShareAlike 4.0 International License.
References
Adamczyk B, Simon J, Kitunen V, Adamczyk S, Smolander A. 2017. Tannins and their complex interaction with different organic nitrogen compounds and enzymes: Old paradigms versus recent advances. Chem Open 6 (5): 610-614. https://doi.org/10.1002/open.201700113.
Al-Khayri JM, Rashmi R, Toppo V, Chole PB, Banadka A, Sudheer WN, Nagella P, Shehata WF, Al-Mssallem MQ, Alessa FM, Almaghasla MI, Rezk AA-S. 2023. Plant secondary metabolites: The weapons for biotic stress management. Metabolites 13 (6): 716. https://doi.org/10.3390/metabo13060716.
Arzate-Vázquez I, Méndez-Méndez JV, Nicolás-Bermúdez J, Chanona-Pérez JJ, Domínguez-Fernández RN, Vélez-Rivera N. 2022. Effect of calcium oxalate crystals on the micromechanical properties of sclerenchyma tissue from the pecan nutshell (Carya illinoinensis). Plant Physiol Biochem 170: 249-254. https://doi.org/10.1016/j.plaphy.2021.12.011.
Azhar B, Gunawan S, Setyadi ERF, Majidah L, Taufany F, Atmaja L, Aparamarta HW. 2023. Purification and separation of glucomannan from porang tuber flour (Amorphophallus muelleri) using microwave assisted extraction as an innovative gelatine substituent. Heliyon 9 (11): e21972. https://doi.org/10.1016/j.heliyon.2023.e21972.
Bele AA, Jadhav VM, Kadam VJ. 2010. Potential of tannnins: A review. Asian J Plant Sci 9 (4): 209-214. https://doi.org/10.3923/ajps.2010.209.214.
Ceretto V, Nacca N. 2018. Mucosal injury from calcium oxalate crystals resembling anaphylaxis and angioedema. J Emerg Med 55 (5): 666-669. https://doi.org/10.1016/j.jemermed.2018.07.016.
Cerritos-Castro IT, Patrón-Soberano A, Bojórquez-Velázquez E, González-Escobar JL, Vargas-Ortiz E, Muñoz-Sandoval E, de la Rosa APB. 2022. Amaranth calcium oxalate crystals are associated with chloroplast structures and proteins. Microsc Res Tech 85 (11): 3694-3706. https://doi.org/10.1002/jemt.24221.
Chairiyah N, Harijati N, Mastuti R. 2011. Calcium oxalate (CaOx) crystals in porang (Amorphopallus muelleri Blume) exposed and unexposed sun. Natural B 1 (2): 130-138. https://doi.org/10.21776/ub.natural-b.2011.001.02.6. [Indonesian]
Ciriminna R, Petri GL, Angellotti G, Fontananova E, Meneguzzo F, Luque R, Pagliaro M. 2025. Tannin: An insight into its cosmeceutical properties and uses. Glob Chall 9 (8): 2500115. https://doi.org/10.1002/gch2.202500115.
Costa ER, Demarco D. 2024. Development and holocrine secretion of resin ducts in Kielmeyera appariciana (Calophyllaceae). Plants 13 (13): 1757. https://doi.org/10.3390/plants13131757.
Dereje B. 2021. Composition, morphology and physicochemical properties of starches derived from indigenous Ethiopian tuber crops: A review. Intl J Biol Macromol 187: 911-921. https://doi.org/10.1016/j.ijbiomac.2021.07.188.
Divekar PA, Narayana S, Divekar BA, Kumar R, Gadratagi BG, Ray A, Singh AK, Rani V, Singh V, Singh AK, Kumar A, Singh RP, Meena RS, Behera TK. 2022. Plant secondary metabolites as defense tools against herbivores for sustainable crop protection. Intl J Mol Sci 23 (5): 2690. https://doi.org/10.3390/ijms23052690.
Dodo. 2018. Evaluation of Rare Plant Reintroduction. Pros Sem Nas Masy Biodiv Indon 4 (2): 280-283. https://doi.org/10.13057/psnmbi/m040232. [Indonesian]
Dogramaci M, Dobry EP, Fortini EA, Sarkar D, Eshel D, Campbell MA. 2024. Physiological and molecular mechanisms associated with potato tuber dormancy. J Exp Bot 75 (19): 6093-6109. https://doi.org/10.1093/jxb/erae182.
Dowling C, Blaesing D. 2022. Plant Sampling for Agriculture - A Guide. Fertilizer Australia, Canberra.
Faristy AFP, Vauzia V. 2024. Population study of the Amorphophallus titanum Becc. in Sumatera. J Biologi Universitas Andalas 12 (2): 79-85. https://doi.org/10.25077/jbioua.12.2.79-85.2024. [Indonesian]
Fraga-Corral M, Otero P, Cassani L, Echave J, Garcia-Oliveira P, Carpena M, Chamorro F, Lourenço-Lopes C, Prieto MA, Simal-Gandara J. 2021. Traditional applications of tannin rich extracts supported by scientific data: Chemical composition, bioavailability and bioaccessibility. Foods 10 (2): 251. https://doi.org/10.3390/foods10020251.
Gaveau DLA, Santos L, Locatelli B, Salim MA, Husnayaen H, Meijaard E, Heatubun C, Sheil D. 2021. Forest loss in Indonesian New Guinea (2001-2019): Trends, drivers and outlook. Biol Conserv 261: 109225. https://doi.org/10.1016/j.biocon.2021.109225.
Handayani TD, Trimedona N, Zebua EA, Utama RS. 2023. The effect of pre-drying soaking on calcium oxalate content of porang flour in Lima Puluh Kota Regency, West Sumatra. J Sains Teknologi Pangan 8 (3): 6352-6359. [Indonesian]
He H, Li D, Li X, Fu L. 2024. Research progress on the formation, function, and impact of calcium oxalate crystals in plants. Crystallogr Rev 30 (1): 31-60. https://doi.org/10.1080/0889311x.2024.2309486.
Islam F, Labib RK, Zehravi M, Lami MS, Das R, Singh LP, Mandhadi JR, Balan P, Khan J, Khan SL, Nainu F, Nafady MH, Rab SO, Emran TB, Wilairatana P. 2023. Genus Amorphophallus: A comprehensive overview on phytochemistry, ethnomedicinal uses, and pharmacological activities. Plants 12 (23): 3945. https://doi.org/10.3390/plants12233945.
Jasim RS. 2018. Antioxidant, antimicrobial activities and phytochemical constituents of Cichorium intybus L. aerial parts. Intl J Bot 14 (1): 24-29. https://doi.org/10.3923/ijb.2018.24.29.
Jaume DF, Pelliza YI, Nanni A, Tadey M. 2025. Drought resistance or herbivory defense strategy? Oxalate druses function in a forage xeric species. Funct Plant Biol 52 (3): FP24299. https://doi.org/10.1071/fp24299.
Kang L, Kaur J, Winkeler K, Kubiak D, Hill JE. 2023. How the volatile organic compounds emitted by corpse plant change through flowering. Sci Rep 13: 372. https://doi.org/10.1038/s41598-022-27108-8.
Karabourniotis G, Horner HT, Bresta P, Nikolopoulos D, Liakopoulos G. 2020. New insights into the functions of carbon-calcium inclusions in plants. New Phytol 228 (3): 845-854. https://doi.org/10.1111/nph.16763.
Khan MI, Pandith SA, Shah MA, Reshi ZA. 2023. Calcium oxalate crystals, the plant ‘gemstones’: Insights into their synthesis and physiological implications in plants. Plant Cell Physiol 64 (10): 1124-1138. https://doi.org/10.1093/pcp/pcad081.
Konyar ST, Öztürk N, Dane F. 2014. Occurrence, types and distribution of calcium oxalate crystals in leaves and stems of some species of poisonous plants. Bot Stud 55: 32. https://doi.org/10.1186/1999-3110-55-32.
Lawrie NS, Cuetos NM, Sini F, Salam GA, Ding H, Vancolen A, Nelson JM, Erkens RHJ, Perversi G. 2023. Systematic review on raphide morphotype calcium oxalate crystals in angiosperms. AoB Plant 15 (4): plad031. https://doi.org/10.1093/aobpla/plad031.
Li C, Chen C, Qin L, Zheng D, Du Q, Hou Q, Wen X. 2023. A highlightedly improved method for isolating and characterizing calcium oxalate crystals from tubercles of Mammillaria schumannii. Plant Method 19: 135. https://doi.org/10.1186/s13007-023-01110-1.
Li L. 2025. Impacts of land-use change on biodiversity of tropical forests. Theor Nat Sci 81 (1): 19-24. https://doi.org/10.54254/2753-8818/2025.19755.
Li P, Liu C, Luo Y, Shi H, Li Q, PinChu C, Li X, Yang J, Fan W. 2022. Oxalate in plants: Metabolism, function, regulation, and application. J Agric Food Chem 70 (51): 16037-16049. https://doi.org/10.1021/acs.jafc.2c04787.
Lindeboom N, Chang PR, Tyler RT. 2004. Analytical, biochemical and physicochemical aspects of starch granule size, with emphasis on small granule starches: A review. Starch 56 (3-4): 89-99. https://doi.org/10.1002/star.200300218.
Liu D, Zhang P, Liu D, Feng Y, Chi M, Guo Z, Wang X, Zhong J, Sun M. 2023. An analysis of volatile compounds and study of release regularity in the flower of Amorphophallus titanum in four periods. Horticulturae 9 (4): 487. https://doi.org/10.3390/horticulturae9040487.
Liu Q, Luo L, Zheng L. 2018. Lignins: Biosynthesis and biological functions in plants. Intl J Mol Sci 19 (2): 335. https://doi.org/10.3390/ijms19020335.
Liu Q, Zhou Y, Fettke J. 2021. Starch granule size and morphology of Arabidopsis thaliana starch-related mutants analyzed during diurnal rhythm and development. Molecules 26 (19): 5859. https://doi.org/10.3390/molecules26195859.
Makiyah A, Tresnayanti S. 2017. Acute toxicity test measured by determining the LD50 of ethanol extract of iles-iles tuber (Amorphophallus variabilis Bl.) in white rats strain Wistar. Bandung Med J 49 (3): 145-155. https://doi.org/10.15395/mkb.v49n3.1130. [Indonesian]
Martínez-Quezada DM, Rojas-Leal A, Villaseñor JL, Terrazas T. 2025. Structural considerations and differences between leaf canals and secretory cavities in Asteraceae. Protoplasma 262: 707-720. https://doi.org/10.1007/s00709-024-02028-8.
Miyamoto M, Noma M, Ishii J, Yoshihara S. 2021. Oral symptoms caused by toxic plants containing calcium oxalate. J Pediatr 230: 258-259. https://doi.org/10.1016/j.jpeds.2020.11.001.
Molnar M, Kovač MJ, Pavić V. 2024. A comprehensive analysis of diversity, structure, biosynthesis and extraction of biologically active tannins from various plant-based materials using deep eutectic solvents. Molecules 29 (11): 2615. https://doi.org/10.3390/molecules29112615.
Musazadeh V, Rostami RY, Moridpour AH, Hosseini ZB, Nikpayam O, Falahatzadeh M, Faghfouri AH. 2024. The effect of glucomannan supplementation on lipid profile in adults: A GRADE-assessed systematic review and meta-analysis. BMC Cardiovasc Disord 24: 545. https://doi.org/10.1186/s12872-024-04223-0.
Paiva EAS. 2019. Are calcium oxalate crystals a dynamic calcium store in plants? New Phytol 223 (4): 1707-1711. https://doi.org/10.1111/nph.15912.
Pambudi AY, Harijati N, Arumingtyas EL. 2020. Physiological and genetic variations of Amorphophallus variabilis in Bojonegoro based on glucomannan content, calcium oxalate and RAPD markers. J Exp Life Sci 10: 49-54. https://doi.org/10.21776/ub.jels.2019.010.01.09.
Pfister B, Zeeman SC. 2016. Formation of starch in plant cells. Cell Mol Life Sci 73: 2781-2807. https://doi.org/10.1007/s00018-016-2250-x.
Pizzi A. 2019. Tannins: Prospectives and actual industrial applications. Biomolecules 9 (8): 344. https://doi.org/10.3390/biom9080344.
Plunkert ML, Martínez-Gómez J, Madrigal Y, Hernández AI, Tribble CM. 2024. Tuber, or not tuber: Molecular and morphological basis of underground storage organ development. Curr Opin Plant Biol 80: 102544. https://doi.org/10.1016/j.pbi.2024.102544.
Pongrac P, Serra TS, Castillo-Michel H, Vogel-Mikuš K, Arčon I, Kelemen M, Jenčič B, Kavčič A, Carvalho MTV, Aarts MGM. 2018. Cadmium associates with oxalate in calcium oxalate crystals and competes with calcium for translocation to stems in the cadmium bioindicator Gomphrena claussenii. Metallomics 10 (11): 1576-1584. https://doi.org/10.1039/c8mt00149a.
Prasad R, Shivay YS. 2017. Oxalic acid/oxalates in plants: From self-defence to phytoremediation. Curr Sci 112 (8): 1665-1667. https://doi.org/10.18520/cs/v112/i08/1665-1667.
Prychid CJ, Jabaily RS, Rudall PJ. 2008. Cellular ultrastructure and crystal development in Amorphophallus (Araceae). Ann Bot 101 (7): 983-995. https://doi.org/10.1093/aob/mcn022.
Rakatama A, Pandit R. 2020. Reviewing social forestry schemes in Indonesia: Opportunities and challenges. For Policy Econ 111: 102052. https://doi.org/10.1016/j.forpol.2019.102052.
Rambey R, Rahmawaty R, Rauf A, Nababan ESM, Delvian D, Aththorick TA, Ismail MH, Saputra MH, Gandaseca S, Suratman MN. 2025. Distribution and habitat suitability of Amorphophallus gigas with MaxEnt modeling in North Sumatra, Indonesia. J Threat Taxa 17 (1): 26370-26384. https://doi.org/10.11609/jott.9022.17.1.26370-26384.
Rashwan AK, Younis HA, Abdelshafy AM, Osman AI, Eletmany MR, Hafouda MA, Chen W. 2024. Plant starch extraction, modification, and green applications: A review. Environ Chem Lett 22: 2483-2530. https://doi.org/10.1007/s10311-024-01753-z.
Ruzin S. 1999. Plant Micro Technique and Microscopy. Oxford University Press, Oxford.
Santosa E, Lian CL, Mine Y, Takahata K, Sugiyama N. 2018. Isolating microsatellite from Amorphophallus variabilis and its application for population study in Dramaga Conservation Forest, Indonesia. Biotropia 25 (1): 22-32. https://doi.org/10.11598/btb.2018.25.1.652.
Santosa E, Sugiyama N, Kawabata S, Hikosaka S. 2012. Genetic variations of Amorphophallus variabilis Blume (Araceae) in Java using AFLP. J Agronomi Indonesia 40 (1): 62-68. https://doi.org/10.24831/jai.v40i1.14941.
Santulli C, Gabrielli S, Roselli G. 2025. Use and roles of tannins in polysaccharide-based bioplastics and biocomposites. Organics 6 (2): 19. https://doi.org/10.3390/org6020019.
Sari AK, Indriyani S, Ekowati G, Batoro J. 2017. Diversity of starch grain structure, starch content, and clustering of eight tuberous plant taxa in Simo Kendal, Ngawi Regency. Biotropika J Trop Biol 5 (1): 14-21. https://doi.org/10.21776/ub.biotropika.2017.005.01.3. [Indonesian]
Sari NY, Putra ETS. 2019. The contribution of calcium to changes in leaf anatomical character of oil palm seedlings (Elaeis guineensis Jacq.) under drought stress. Agric Sci 4 (1): 23-32. https://doi.org/10.22146/ipas.42447.
Setiawan RB, Yusniwati Y, Wati DR, Sari IP, Rahmah S, Haryanti S, Bosma PAL. 2025. Conservation comprehensive approach: Study on exploration, habitat analysis, propagation, and reintroduction of the indonesian endemic endangered titan arum (Amorphophallus titanum Becc.). J Trop For Manag 31 (3): 261-273. https://doi.org/10.7226/jtfm.31.3.261.
Shi H-D, Zhang W-Q, Lu H-Y, Zhang W-Q, Ye H, Liu D-D. 2020. Functional characterization of a starch synthesis-related gene AmAGP in Amorphophallus muelleri. Plant Signal Behav 15 (11): 1805903. https://doi.org/10.1080/15592324.2020.1805903.
Simon TK, Nayagam JR. 2018. Ergastic crystal studies for raw drug analysis. In: Builders PF (eds). Herbal Medicine. Intech Open, London. https://doi.org/10.5772/intechopen.74278.
Siroka Z. 2023. Toxicity of house plants to pet animals. Toxins 15 (5): 346. https://doi.org/10.3390/toxins15050346.
Sood N, Baker WL, Coleman CI. 2008. Effect of glucomannan on plasma lipid and glucose concentrations, body weight, and blood pressure: Systematic review and meta-analysis. Am J Clin Nutr 88 (4): 1167-1175. https://doi.org/10.1093/ajcn/88.4.1167.
Susmianto A. 2017. Restoring the love of the King in Gunung Leuser National Park: “My home is starting to stand again - Thank you, uncle Keleng”). In: Rudianto W, Rismayani R, Sulistiyo RA, Adi SW, Chandra A, Fauzi D, Khatimah FH (eds.). Success Stories of Participatory Ecosystem Restoration in Nature Reserves and Nature Conservation Areas. Forda Press, Jakarta. [Indonesian]
Tooulakou G, Giannopoulos A, Nikolopoulos D, Bresta P, Dotsika E, Orkoula MG, Kontoyannis CG, Fasseas C, Liakopoulos G, Klapa MI, Karabourniotis G. 2016. Alarm photosynthesis: Calcium oxalate crystals as an internal CO2 source in plants. Plant Physiol 171 (4): 2577-2585. https://doi.org/10.1104/pp.16.00111.
Victório CP, dos Santos MS, Dias AC, Bento JPSP, dos Santos FBH, da Costa SM, Simas NK, de Oliveira ARdC. 2023. Laguncularia racemosa leaves indicate the presence of potentially toxic elements in mangroves. Sci Rep 13: 4845. https://doi.org/10.1038/s41598-023-31986-x.
Wahidah BF, Afiati N, Jumari. 2022. Ecological role and potential extinction of Amorphophallus variabilis in Central Java, Indonesia. Biodiversitas 23 (4): 1765-1773. https://doi.org/10.13057/biodiv/d230407.
War AR, Paulraj MG, Ahmad T, Buhroo AA, Hussain B, Ignacimuthu S, Sharma HC. 2012. Mechanisms of plant defense against insect herbivores. Plant Signal Behav 7 (10): 1306-1320. https://doi.org/10.4161/psb.21663.
Wardani RK, Arifiyana D. 2020. The Effect of Soaking Time and Temperature of Acetic Acid Solution to the Decrease of Calcium Oxalate Levels in Porang Tubers. 1st Intl Conf Eco-Innov Sci Eng Technol. NST Proc. UPN Veteran Jawa Timur, Surabaya, 28 September 2020. https://doi.org/10.11594/nstp.2020.0522.
Weber E, Levy D, Sasson MB, Fitch AN, Pokroy B. 2015. Structural analysis of metal-doped calcium oxalate. RSC Adv 5 (119): 98626-98633. https://doi.org/10.1039/c5ra20870j.
Xie X, Chen J, Cheng L, Zhang B, Zhu H, Xu C, Liang D. 2024. Physicochemical properties of different size fractions of potato starch cultivated in Highland China. Intl J Biol Macromol 256 (Part 1): 128065. https://doi.org/10.1016/j.ijbiomac.2023.128065.
Yu T, Yang Q, Deng M, Cheng N, Yao K, Yang W, Ji X, Zheng H. 2022. Morphological characteristics of seed starch granules of Fagaceae in South China and their implication in paleodiet. Front Plant Sci 13: 977152. https://doi.org/10.3389/fpls.2022.977152.
Zavala-García LE, Sánchez-Segura L, de Dios EA, Pérez-López A, Simpson J. 2018. Starch accumulation is associated with active growth in A. tequilana. Plant Physiol Biochem 130: 623-632. https://doi.org/10.1016/j.plaphy.2018.08.011.