PERTANIKA JOURNAL OF TROPICAL AGRICULTURAL SCIENCE

The Garcinia genus belongs to the Clusiaceae or Guttiferae family, comprising 350 species. Garcinia is a large genus, and it is widely distributed in tropical environments, especially in the Southeast Asia region. Despite their wide distribution in Malaysia, information on their reproductive biology is still lacking. This overview highlights the distribution and the reproductive system of several Garcinia species in Malaysia, which are Garcinia mangostana var. mangostana, Garcinia celebica, Garcinia mangostana var. malaccensis, Garcinia prainiana, Garcinia cowa, Garcinia atroviridis and Garcinia parvifolia. Apomixis, specifically agamospermy in Garcinia species, is widely acknowledged by previous research. Apomixis develops a distinctive mechanism in gametophytic and sporophytic types, and it is molecularly triggered either by hybridisation or polyploidisation. G. mangostana var. mangostana is classified under obligate apomict due to male sterility caused by the alteration of tapetum during pollen development. On the other hand, the occurrence of male trees, male fertility and pollen viability are the important features that consider G. celebica, G. mangostana var. malaccensis, G. prainiana, G. cowa, G. atroviridis and G. parvifolia to be facultative apomict. Besides, the seeds of Garcinia species are recalcitrant and possess low resistance to desiccation. The size and moisture content of the seeds can influence germination ability. The overall information of this review can help explore the life cycle of Garcinia trees to facilitate conservation programs along with the agricultural benefits.

• Ahmar, S., Gill, R. A., Jung, K. H., Faheem, A., Qasim, M. U., Mubeen, M., & Zhou, W. (2020). Conventional and molecular techniques from simple breeding to speed breeding in crop plants: Recent advances and future outlook. International Journal of Molecular Sciences, 21(7), 1-24. http://doi.org/10.3390/ijms21072590
  
  Ao, C. (2020). The origin of double fertilization in flowering plants: Looking into the history of plant reproduction evolution. Plant Biosystems - An International Journal Dealing with All Aspects of Plant Biology, 155(5), 994–1000. https://doi.org/10.1080/11263504.2020.1810813
  
  Apiratikorn, S., Sdoodee, S., Lerslerwong, L., & Rongsawat, S. (2012). The impact of climatic variability on phenological change, yield and fruit quality of mangosteen in Phatthalung Province, Southern Thailand. Agriculture and Natural Resources, 46(1), 1-9.
  
  Ardiarini, N., Lase, J., Hidayat, Y., & Habeahan, K. (2021). The effect of seed scarification on the germination process and the growth of long bean (Vigna sinensis) sprout. E3S Web of Conferences, 306(01002). https://doi.org/10.1051/e3sconf/202130601002
  
  Awachare, C. M., & Upreti, K. K. (2019). Phenological growth stages in mangosteen (Garcinia mangostana L.) according to the extended BBCH scale. Annals of Applied Biology, 176(1), 16-25. https://doi.org/10.1111/aab.12552
  
  Azuan, A., Salma, I., Mirfat, A. H. S., Erny, S. M. N., Mohd Saifuddin, M., & Rusli, A. (2015). Diversity of Garcinia prainiana: An underutilized fruit species. In Proceeding Regional Taxonomy and Ecology Conference 2015. https://www.researchgate.net/publication/328332702_Diversity_Of_Garcinia_Prainiana_An_Underutilized_Fruit_Species
  
  Barke, B. H., Daubert, M., & Hörandl, E. (2018). Establishment of apomixis in diploid F2 hybrids and inheritance of apospory from F1 to F2 hybrids of the Ranunculus auricomus complex. Frontiers in Plant Science, 9, 1111. https://doi.org/10.3389/fpls.2018.01111
  
  Bayu, E. S., Febrianti, W., & Damanik, R. I. M. (2018). Flower morphology diversity of asam gelugur (Garcinia atroviridis Griff. ex T. Anders) accessions in several districts of North Sumatera, Indonesian. IOP conference series: Earth and environmental science, 122, 012038. https://doi.org/10.1088/1755-1315/122/1/012038
  
  Bayu, E. S., Lestami, A., Kardhinata, E., & Rosemary (2018). Phylogenetic relationship of asam gelugur (Garcinia atroviridis Griff. ex T. Anders) based on morphological characters in Langkat and Serdang Berdagai, Sumatera Utara. IOP conference series: Earth and environmental science, 122, 012077. https://doi.org/10.1088/1755-1315/122/1/012077
  
  Berame, J., Lawsin, N., Migue, F., & Chavez, J. (2020). Morphological variations of mangosteen fruits from Luzon and Mindanao Islands, the Philippines. Biodiversitas, 21(7), 3094-3100. https://doi.org/10.13057/biodiv/d210729
  
  Bitew, Y., & Alemayahu, M. (2017). Impact of crop production inputs on soil health: A review. Asian Journal of Plant Sciences, 16(3), 109-131. https://doi.org/10.3923/ajps.2017.109.131
  
  Brukhin, V. (2017). Molecular and genetic regulation of apomixis. Russian Journal of Genetics, 53, 943–964. https://doi.org/10.1134/S1022795417090046
  
  Cardoso, J. C. F., Viana, M. L., Matias, R., Furtado, M. T., Caetano, A. P. S., Consolaro, H., & Brito, V. L. G. (2018). Towards a unified terminology for angiosperm reproductive systems. Acta Botanica Brasilica, 32(3), 329–348. https://doi.org/10.1590/0102-33062018abb0124
  
  Cruz, F. S. D. J. (2001). Status report on genetic resources of Mangosteen (Garcinia mangostana L.) in Southeast Asia. IPGRI Office for South Asia. https://scholar.google.com/scholar_lookup?title=Status report on genetic resources of Mangosteen (Garcinia mangostana L.) in Southeast Asia&author=F. S. D. J. Cruz&publication_year=2001&
  
  Debeaujon, I., Léon-Kloosterziel, K. M., & Koornneef, M. (2000). Influence of the testa on seed dormancy, germination, and longevity in Arabidopsis. Plant physiology, 122(2), 403–414. https://doi.org/10.1104/pp.122.2.403
  
  Deepu, S., & Geethakumary, M. P. (2020, August 20). Cowa fruit. IUCN red list. https://doi.org/10.2305/IUCN.UK.2020-3.RLTS.T61988928A61988973.en
  
  den Nijs, A. P. M., & van Dijk, G. E. (1993). Apomixis. In M. D. Hayward, N. D. Bosemark, & I. Romagosa (Eds.), Plant breeding: Principles and prospect. Chapman and Hall.
  
  Dike, M. S., Malik, S. K., Sawardekar, S. V., & Deodhar, M. A. (2020). Study of the mode of reproduction and fruit development in Garcinia Indica. International Journal of Fruit Science, 20(1), 20-38. https://doi.org/10.1080/15538362.2018.1563022
  
  Goh, H. H., Syuhaidah, A. B., Nur, D. K. A., Zamri, Z., & Normah, M. N. (2019). Transcriptional reprogramming during Garcinia-type recalcitrant seed germination of Garcinia mangostana. Scientia Horticulturae, 257, 108727. https://doi.org/10.1016/j.scienta.2019.108727
  
  Ha, C. O., Sands, V. E., Soepadmo, E., & Jong, K. (1988). Reproductive patterns of selected understorey trees in the Malaysian rain forest: The apomictic species. Botanical Journal of the Linnean Society, 97(3), 317-331. https://doi.org/10.1111/j.1095-8339.1988.tb01586.x
  
  Hand, M. L., & Koltunow, A. M. (2014). The genetic control of apomixis: Asexual seed formation. Genetics, 197(2), 441–450. https://doi.org/10.1534/genetics.114.163105
  
  Hassan, S. H. A., Fry, J. R., & Abu Bakar, M. F. (2013). Phytochemicals content, antioxidant activity and acetylcholinesterase inhibition properties of indigenous Garcinia parvifolia fruit. BioMed Research International, 2013(1), 138950. http://doi.org/10.1155/2013/138950
  
  Hojsgaard, D., & Hörandl, E. (2018). The rise of apomixis in natural plant populations. Frontiers in Plant Science, 10(358). https://doi.org/10.3389/fpls.2019.00358
  
  Idris, S., & Rukayah, A. (1987). Description of male mangosteen (Garcinia mangostana L.) discovered in Peninsular Malaysia. Mardi Research Bulletin, 15(1), 63–66.
  
  John, K. J., Kumar, R. S., Suresh, C. P., George, J. K., & Abraham, Z. (2008). Occurrence, distribution and economic potential of seashore mangosteen (Garcinia hombroniana Pierre) in India. Genetic Resources Crop and Evolution, 55, 183-186.
  
  Kant, R., & Verma, J. (2012). Obligate apomixis in Zeuxine strateumatica (Lindl.) Schltr. (Orchidaceae). International Journal of Plant Research, 25(1), 274-277.
  
  Khoo, H. E., Prasad, K., Kong, K. W., Chew, L.Y., Azlan, A., Sun, J., Ismail, A., & Idris, S. (2010). A review on underutilized tropical fruits in Malaysia. Guangxi Agricultural Sciences, 41(7), 698-702.
  
  Koltunow, A. M., Bicknell, R. A., & Chaudhury, A. M. (1995). Apomixis: Molecular strategies for the generation of genetically identical seeds without fertilization. Plant Physiology, 108(4), 1345-1352. https://doi.org/10.1104/pp.108.4.1345
  
  Li, T., Liu, X., Li, Z., Ma, H., Wan, Y., Liu, X., & Fu, L. (2018). Study on reproductive biology of Rhododendron longipedicellatum: A newly discovered and special threatened plant surviving in limestone habitat in Southeast Yunnan, China. Frontiers in Plant Science, 9, 33. https://doi.org/10.3389/fpls.2018.00033
  
  Lim, A. L. (1984). The embryology of Garcinia mangostana L. (Clusiaceae). Gardens' Bulletin Singapore, 37(1), 93–103.
  
  Lim, T. K. (2012a). Garcinia mangostana. In Edible medicinal and non-medicinal plants (Vol. 2, pp. 83–108). Springer. https://doi.org/10.1007/978-94-007-1764-0_15
  
  Lim, T. K. (2012b). Garcinia hombroniana. In Edible medicinal and non-medicinal plants (Vol. 2, pp. 56-58). Springer. https://doi.org/10.1007/978-94-007-1764-0_8
  
  Lim, T. K. (2012c). Garcinia malaccensis. In Edible medicinal and non-medicinal plants (Vol. 2, pp. 80-82). Springer. https://doi.org/10.1007/978-94-007-1764-0_14
  
  Lim, T. K. (2012d). Garcinia prainiana. In Edible medicinal and non-medicinal plants (Vol. 2, pp. 120-122). Springer. https://doi.org/10.1007/978-94-007-1764-0_19
  
  Lim, T. K. (2012e). Garcinia cowa. In Edible medicinal and non-medicinal plants (Vol. 2, pp. 29-34). Springer. https://doi.org/110.1007/978-94-007-1764-0_4
  
  Lim, T. K. (2012f). Garcinia atroviridis. In Edible medicinal and non-medicinal plants (Vol. 2, pp. 21-28). Springer. https://doi.org/10.1007/978-94-007-1764-0_3
  
  Lim, T. K. (2012g). Garcinia parvifolia. In Edible medicinal and non-medicinal plants (Vol. 2, pp. 115-119). Springer. https://doi.org/10.1007/978-94-007-1764-0_18
  
  Liu, Y., Chen, J., Bai, Z. L., Deng, X. B., & Zhang, L. (2002). Seed dispersal, seed predation, and seedling spatial pattern of Garcinia cowa (Guttiferae). Chinese Journal of Plant Ecology, 26(4), 427–434.
  
  Liu, Y., Qiu, Y. P., Zhang, L., & Chen, Jin. (2005). Dormancy breaking and storage behaviour of Garcinia cowa Roxb. (Guttiferae) seeds: Implications for ecological function and germplasm conservation. Journal of Integrative Plant Biology, 47(1). https://doi.org/10.1111/j.1744-7909.2005.00010.x
  
  Majeský, L., Krahulec, F., & Vašut, R. J. (2017). How apomictic taxa are treated in current taxonomy: A review. Taxon, 66(5), 1017-1040.
  
  Mansyah, E., Baihaki, A., Setiamihardja, R., Darsa, J. S., Sobir, & Poerwanto, R. (2003). Phenotypic variability of mangosteen (Garcinia mangostana L.) at several production centres in Java. Jurnal Hortikultura, 13, 147-156.
  
  Mansyah, E., Muas, I., & Jawal, M. (2010). Morphological variability of apomictic mangosteen (Garcinia mangostana L.) in Indonesia: Morphological evidence of natural populations from Sumatra and Java. SABRAO Journal of Breeding and Genetics, 42(1), 1-8.
  
  Marbaniang, E. J., Venugopal, N., Verma, S., Raina, R., Khajuria, A., & Gautam, K. (2018). Floral biology and embryological studies are important for conservation of threatened plants having reproductive bottlenecks: A case study of Illicium griffithii Hook. f. & Thomson. Current Science, 114(3), 576-587. https://doi.org/10.18520/cs/v114/i03/576-587
  
  Martias, Sparta, A., Hendri, Fatria, D., Marlina, L., & Mansyah, E. (2021). The potency of sub-micron dolomite in reducing yellow latex contamination in mangosteen fruits. IOP conference series: Earth and environmental science, 648, 012139. https://doi.org/10.1088/1755-1315/648/1/012139
  
  Mulyono, D., Irawati, Y., & Syah, M. J. A. (2021). Identification morphological variability of six mangosteen (Garcinia mangostana L.) as a conservation strategy for local varieties. IOP conference series: Earth and environmental science, 739, 012076. https://doi.org/10.1088/1755-1315/739/1/012076
  
  Nazre, M. (2010). Historical review and notes on the correct scientific name for seashore mangosteen. Genetic Resources and Crop Evolution, 57, 1249–1259. https://doi.org/10.1007/s10722-010-9588-y
  
  Nazre, M. (2014). New evidence on the origin of mangosteen (Garcinia mangostana L.) based on morphology and ITS sequence. Genetic Resources and Crop Evolution, 61(6). https://doi.org/10.1007/s10722-014-0097-2
  
  Nazre, M., Newman, M. F., Pennington, R. T. & Middleton, D. J. (2018). Taxonomic revision of Garcinia section Garcinia (Clusiaceae). Phytotaxa, 373(1). https://doi.org/10.11646/phytotaxa.373.1.1
  
  Normah, M. N., Wan, M. A., Khatijah, H., & Emelda, R. R. (2016). Seed characteristics and germination properties of four Garcinia (Clusiaceae) fruit species. Fruits, 71(4), 99–207. https://doi.org/10.1051/fruits/2016008
  
  Nuanjunkong, N., & Meesawat, U. (2010, October 7-8). Anatomical changes during pollen development of apomictic mangosteen (Garcinia mangostana L.) [Paper Presentation]. Proceeding The 7th IMT-GT Uninet and the 3rd Joint International PSU-UNS Conferences, Prince of Songkla University, Hat Yai, Songkhla, Thailand. https://www.researchgate.net/publication/273441708_Anatomical_Changes_During_Pollen_Development_of_Apomictic_Mangosteen_Garcinia_mangostana_L
  
  Mohd Khairuddin, N. N., Gogula, S. A., Ahamd, A., Wan, R. W. T., & Alias, N. (2018). Evaluation of antioxidant, antimicrobial activity and phytochemical content of Garcinia prainiana-An endangered plant. Research Journal of Pharmacy and Technology, 11(9), 3752-3758. https://doi.org/10.5958/0974-360X.2018.00687.X
  
  Oliveira, L. M. Q., & Valio, I. F. M. (1992). Effects of moisture content on germination of seeds of Hancornia speciosa Gom. (Apocynaceae). Annals of Botany, 69(1), 1–5. https://doi.org/10.1093/oxfordjournals.aob.a088299
  
  Osman, M., & Milan, A. R. (2006). Mangosteen: Garcinia mangostana L. University of Southampton. http://eprints.soton.ac.uk/id/eprint/53412
  
  Pangsuban, S., Bamroongrugsa, N., Kanchanapoom, K., & Nualsri, C. (2009). Facultative apomixis in Garcinia atroviridis (Clusiaceae) and effects of different pollination regimes on reproductive success. Tropical Life Sciences Research, 20(2), 89–108.
  
  Pareek, A., Dhankher, O. P., & Foyer, C. H. (2020). Mitigating the impact of climate change on plant productivity and ecosystem sustainability. Journal of Experimental Botany, 71(2), 451–456. https://doi.org/10.1093/jxb/erz518
  
  Puixeu, G., Pickup, M., Field, D. L., & Barrett, S. C. H. (2019). Variation in sexual dimorphism in a wind-pollinated plant: The influence of geographical context and life-cycle dynamics. New Phytologist, 224(3), 1108–1120. https://doi.org/10.1111/nph.16050
  
  Richards, A. J. (1990a). Studies in Garcinia, dioecious tropical fruit trees: Agamospermy. Botanical Journal of the Linnean Society, 103, 233-250. https://doi.org/10.1111/j.1095-8339.1990.tb00186.x
  
  Richards, A. J. (1990b). Studies in Garcinia, dioecious tropical fruit trees: The origin of the mangosteen (G. mangostana L.). Botanical Journal of the Linnean Society, 103, 301–308. https://doi.org/10.1111/j.1095-8339.1990.tb00191.x
  
  Richards, A. J. (1990c). Studies in Garcinia, dioecious tropical forest trees: The phenology, pollination biology and fertilization of G. hombroniana Pierre. Botanical Journal of the Linnean Society, 103, 251-261. https://doi.org/10.1111/j.1095-8339.1990.tb00187.x
  
  Rohani, E. R., Norfadzilah, A. F., Clyde, M. M., & Noor, N. M. (2021). Insights into the mode of reproduction of Garcinia prainiana. Fruits, 76, 22-29. https://doi.org/10.17660/th2021/76.1.3
  
  Roy, D. K., Fakir, M. S. A., Rahman, M. M., & Rahman, M. (2010). Fruit growth in cowphal (Garcinia cowa). Journal of Agroforestry and Environment, 3(2), 52-59.
  
  Shi, S., Ding, D., Mei, S., & Wang, J. (2010). A comparative light and electron microscopic analysis of microspore and tapetum development in fertile and cytoplasmic male sterile radish. Protoplasma, 241, 37–49. https://doi.org/10.1007/s00709-009-0100-5
  
  Sobir, & Poerwanto, R. (2007). Mangosteen genetics and improvement. International Journal of Plant Breeding, 1(2), 105-111.
  
  Steiner, F., Zuffo, A. M., Busch, A., Sousa, T. D. O., & Zoz, T. (2019). Does seed size affect the germination rate and seedling growth of peanut under salinity and water stress? Pesquisa Agropecuaria Tropical, 49, 1-9. https://doi.org/10.1590/1983-40632019v4954353
  
  Suhendra, D., & Mustamu, N. E. (2018). Structure and germination pattern mangosteen seed (Garcinia mangostana L). IOP conference series: Earth and Environmental Science, 205, 012015. https://doi.org/10.1088/1755-1315/205/1/012015
  
  Sulassih, Sobir, & Santosa, E. (2013). Phylogenetic analysis of mangosteen (Garcinia mangostana L.) and its relatives based on morphological and inter simple sequence repeat (ISSR) markers. SABRAO Journal of Breeding and Genetics, 45(3), 478-490.
  
  Sutthinon, P., Samuels, L., & Meesawat, U. (2018a, December 11-12). Insoluble carbohydrate distribution during pollen abortion in mangosteen (Garcinia mangostana L.) compared with pollen development in seashore mangosteen (Garcinia celebica L.) [Paper presentation]. Proceeding of the 11th IMT-GT UNINET Conference 2018 – Bioscience for a Sustainable Future, Penang, Malaysia. https://www.researchgate.net/publication/331167031_Insoluble_Carbohydrate_Distribution_during_Pollen_Abortion_in_Mangosteen_Garcinia_mangostana_L_Compared_with_Pollen_Development_in_Seashore_Mangosteen_Garcinia_celebica_L
  
  Sutthinon, P., Samuels, A. L., & Meesawat, U. (2018b). Male functionality in Garcinia celebica L.: A candidate ancestor species of mangosteen (G. mangostana L.). Botany, 96(2). https://doi.org/10.1139/cjb-2018-0079
  
  Sutthinon, P., Samuels, L., & Meesawat, U. (2019). Pollen development in male sterile mangosteen (Garcinia mangostana L.) and male fertile seashore mangosteen (Garcinia celebica L.). Protoplasma, 256(6), 1545-1556. https://doi.org/10.1007/s00709-019-01397-9
  
  Sweeney, P. W. (2008). Phylogeny and floral diversity in the genus Garcinia (Clusiaceae) and relatives. International Journal of Plant Sciences, 169(9), 1288–1303. https://doi.org/10.1086/591990
  
  Syahputra, M. R., Setiado, H., Siregar, L. A. M., & Damanik, R. I. (2021). Morphological characteristics of mangosteen plants (Garcinia mangostana L.) in Langkat District, North Sumatera, Indonesia. IOP conference series: Earth and Environmental Science. 782, 042056. https://doi.org/10.1088/1755-1315/782/4/042056
  
  Taher, M., Susanti, D., Rezali, M. F., Zohri, F. S., Ichwan, S. J., Alkhamaiseh, S. I., & Ahmad, F. (2012). Apoptosis, antimicrobial and antioxidant activities of phytochemicals from Garcinia malaccensis Hk.f. Asian Pacific Journal of Tropical Medicine, 5(2), 136-141. https://doi.org/10.1016/S1995-7645(12)60012-1
  
  Te-chato, S. (2007). Floral and fruit morphology of some species in Garcinia spp. Songklanakarin Journal of Science and Technology, 29(2), 245-252.
  
  Vijverberg, K., Ozias-Akins, P., & Schranz, M. E. (2019). Identifying and engineering genes for parthenogenesis in plants. Frontiers in Plant Science, 10(128). https://doi.org/10.3389/fpls.2019.00128
  
  Vogel, E. F. (1980). Seedlings of dicotyledons. Feddes Reepertorium, 92(9-10), 712-721. https://doi.org/10.1002/fedr.19810920909
  
  Wen, B., & Cai, Y. (2014). Seed viability as a function of moisture and temperature in the recalcitrant rainforest species Baccaurea ramiflora (Euphorbiaceae). Annals of Forest Science, 71(8), 853-861. https://doi.org/10.1007/s13595-014-0388-y
  
  Whitmore, T. C. (1973). Guttiferae. In T. C. Whitmore (Ed.), Tree flora of Malaya 2 (pp. 162- 236). Longman Malaysia.
  
  Yaacob, O., & Tindall, H. D. (1995). Mangosteen cultivation. Food and Agriculture Organization. https://doi.org/10.5962/bhl.title.121365
  
  Yapwattanaphun, C., Kobayashi, S., Yonemori, K., & Ueda, J. (2014). Hormone analysis in the locule of mangosteen fruit during apomictic seed development. Acta Horticulturae, 1024, 141–146. https://doi.org/10.17660/ActaHortic.2014.1024.15
  
  Yapwattanaphun, C., Tachibana, K., & Yonemori, K. (2008). Pollen abortion in the flower of mangosteen. Acta Horticulturae, 787, 245-250. https://doi.org/10.17660/ActaHortic.2008.787.25
  
  Yousif, A. A. B. (2010). Effect of seed age, size and moisture content on seed quality of sorghum (Sorghum bicolor. L. Moench). Research Journal of Agriculture and Biological Sciences, 6(4), 522-529.
  
  Yuantao, X., Huihui, J., Xiaomeng, W., Koltunow, A. M. G., Xiuxin, D., & Qiang, X. (2021). Regulation of nucellar embryony, a mode of sporophytic apomixis in Citrus resembling somatic embryogenesis. Current Opinion in Plant Biology, 59, 101984. https://doi.org/10.1016/j.pbi.2020.101984
  
  Yuniastuti, E. (2010). Morphological description of Jogorogo Mangosteen (Garcinia mangostana L.). Journal of Biotechnology and Biodiversity, 1(1), 20-25.