Interactive Effects of Fertilizer Types and Shading Levels on Growth, Yield, and Piperine Content of Piper Retrofractum Under Tropical Agroecosystems
Abstract
Piper retrofractum (Javanese long pepper) is an economically important medicinal plant valued for its piperine content. However, its productivity and quality remain inconsistent under tropical agroecosystems due to suboptimal nutrient management and light conditions. This study aimed to evaluate the interactive effects of fertilizer types and shading levels on growth, yield, and piperine content, as well as to determine the relationships among these variables. A factorial completely randomized design was employed with two factors: fertilizer types (NPK, biofertilizer, local microbial liquid organic fertilizer, and a combination of NPK and biofertilizer) and shading levels (0%, 40%, and 60%), resulting in 12 treatment combinations with three replications. The observed parameters included number of productive branches, dry fruit weight per plant, and piperine content. Data were analyzed using ANOVA, followed by DMRT at 5%, and strengthened with Pearson correlation and multiple regression analyses. The results showed that the combination of NPK and biofertilizer under no shading significantly produced the highest values for all parameters. Productive branches exhibited a very strong positive correlation with dry fruit weight (r = 0.88**) and moderate correlation with piperine content (r = 0.61*), while dry fruit weight showed a strong correlation with piperine content (r = 0.69**). The regression model (R² = 0.88) indicated that both productive branches and dry fruit weight significantly contributed to piperine content, with dry fruit weight having a greater influence. These findings highlight that optimizing fertilization strategies and light conditions can simultaneously enhance yield and bioactive compound accumulation in P. retrofractum under tropical conditions.
Downloads
References
Ahmed, S., Roro, A. G., Beshir, H. M., & Haile, A. (2023). Physiological, growth, yield, and quality responses of hot pepper due to shade level. International Journal of Vegetable Science, 29(5), 375–402. https://doi.org/10.1080/19315260.2023.2233955
Ali, A., Niu, G., Masabni, J., Ferrante, A., & Cocetta, G. (2024). Integrated nutrient management of fruits, vegetables, and crops through the use of biostimulants, soilless cultivation, and traditional and modern approaches—A mini review. Agriculture, 14(8), 1330. https://doi.org/10.3390/agriculture14081330
Borbély, P., Gasperl, A., Pálmai, T., Ahres, M., Asghar, M. A., Galiba, G., Müller, M., & Kocsy, G. (2022). Light intensity- and spectrum-dependent redox regulation of plant metabolism. Antioxidants, 11(7), 1311. https://doi.org/10.3390/antiox11071311
Carsono, N., Tumilaar, S. G., Kurnia, D., Latipudin, D., & Satari, M. H. (2022). A review of bioactive compounds and antioxidant activity properties of Piper species. Molecules, 27(19), 6774. https://doi.org/10.3390/molecules27196774
Chong, W.-H., Chin, S.-F., Pang, S.-C., & Kok, K.-Y. (2020). Synthesis and characterisation of piperine-loaded starch nanoparticles. Journal of Physical Science, 31(1), 57–68. https://doi.org/10.21315/jps2020.31.1.4
Cioć, M., Szopa, A., Prokopiuk, B., Pawłowska, B., & Łopusiewicz, Ł. (2025). Influence of LED light spectra on morphogenesis, secondary metabolite production and antioxidant potential in Eucomis autumnalis cultured in vitro. Agronomy, 15(9), 2197. https://doi.org/10.3390/agronomy15092197
Costa, E. F., Magalhães, W. V., & Di Stasi, L. C. (2022). Recent advances in herbal-derived products with skin anti-aging properties and cosmetic applications. Molecules, 27(21), 7518. https://doi.org/10.3390/molecules27217518
Díaz-Guerrero, P., Panzani, S., Sanmartin, C., Muntoni, C., Taglieri, I., & Venturi, F. (2025). “Pepper”: Different spices, one name—Analysis of sensory and biological aspects. Molecules, 30(9), 1891. https://doi.org/10.3390/molecules30091891
Gao, P., Nasution, A. K., Ono, N., Kanaya, S., & Altaf-Ul-Amin, M. (2025). Investigating potential anti-bacterial natural products based on Ayurvedic formulae using supervised network analysis and machine learning approaches. Pharmaceuticals, 18(2), 192. https://doi.org/10.3390/ph18020192
Gelaye, Y. (2025). Intercropping of pepper (Capsicum annuum L.) and black cumin (Nigella sativa L.) optimize crop performance and system productivity in Ethiopia: Systematic review. Cogent Food & Agriculture, 11(1), 2451057. https://doi.org/10.1080/23311932.2025.2451057
Gokkus, M. K. (2025). Effects of urea fertilizer and nettle extract on the biochemical and morphological characteristics of ornamental peppers (Capsicum frutescens L.) under deficit irrigation conditions. Irrigation and Drainage, 74(2), 600–614. https://doi.org/10.1002/ird.3049
Guo, Y., Zhao, C., Liu, X., Dong, Y., Liu, W., Chen, Q., Ding, S., Zhang, J., Guo, B., & Gao, X. (2025). The impact of organic fertilizer substitution on microbial community structure, greenhouse gas emissions, and enzyme activity in soils with different cultivation durations. Sustainability, 17(10), 4541. https://doi.org/10.3390/su17104541
Ibáñez, A., Garrido-Chamorro, S., Vasco-Cárdenas, M., & Barreiro, C. (2023). From lab to field: Biofertilizers in the 21st century. Horticulturae, 9(12), 1306. https://doi.org/10.3390/horticulturae9121306
Izadi, Z., Biabani, A., Sabouri, H., & Bahreininejad, B. (2022). The effect of different levels of urea and planting density on the phytochemical characteristics, alkaloids, and yield of the medicinal plant jimsonweed (Datura stramonium L.). Crop Science, 62(3), 1264–1276. https://doi.org/10.1002/csc2.20733
Jahan, N., Sarker, U., Hasan Saikat, M. M., Hossain, M. M., Azam, M. G., Ali, D., Ercisli, S., & Golokhvast, K. S. (2023). Evaluation of yield attributes and bioactive phytochemicals of twenty amaranth genotypes of Bengal floodplain. Heliyon, 9(9), e19644. https://doi.org/10.1016/j.heliyon.2023.e19644
KC, S., Long, L., Zhang, Q., Ni, K., Ma, L., & Ruan, J. (2022). Effect of interactions between phosphorus and light intensity on metabolite compositions in tea cultivar Longjing43. International Journal of Molecular Sciences, 23(23), 15194. https://doi.org/10.3390/ijms232315194
Khan, A. Q., Guo, C., Ullah, R., Ahmad, F., Li, Z., Ma, H., Feng, T., & Li, F.-M. (2025). Enhancing soil health and microbial resilience with organic fertilization in semi-arid Astragalus ecosystems. Journal of Environmental Management, 394, 127474. https://doi.org/10.1016/j.jenvman.2025.127474
Kumar, S., Diksha, Sindhu, S. S., & Kumar, R. (2022). Biofertilizers: An ecofriendly technology for nutrient recycling and environmental sustainability. Current Research in Microbial Sciences, 3, 100094. https://doi.org/10.1016/j.crmicr.2021.100094
Li, L., Chen, F., & Xing, G. (2022). Effects of fertilizer level and intercropping planting pattern with corn on the yield-related traits and insect community of soybean. Agronomy, 12(12), 3080. https://doi.org/10.3390/agronomy12123080
Li, Y., Kong, D., Fu, Y., Sussman, M. R., & Wu, H. (2020). The effect of developmental and environmental factors on secondary metabolites in medicinal plants. Plant Physiology and Biochemistry, 148, 80–89. https://doi.org/10.1016/j.plaphy.2020.01.006
Liu, Y., Singh, S. K., Pattanaik, S., Wang, H., & Yuan, L. (2023). Light regulation of the biosynthesis of phenolics, terpenoids, and alkaloids in plants. Communications Biology, 6(1), 1055. https://doi.org/10.1038/s42003-023-05435-4
Lu, D., Liu, B., Ren, M., Wu, C., Ma, J., & Shen, Y. (2021). Light deficiency inhibits growth by affecting photosynthesis efficiency as well as JA and ethylene signaling in endangered plant Magnolia sinostellata. Plants, 10(11), 2261. https://doi.org/10.3390/plants10112261
Lv, J., Yang, S., Zhou, W., Liu, Z., Tan, J., & Wei, M. (2024). Microbial regulation of plant secondary metabolites: Impact, mechanisms and prospects. Microbiological Research, 283, 127688. https://doi.org/10.1016/j.micres.2024.127688
Qaderi, M. M., Martel, A. B., & Strugnell, C. A. (2023). Environmental factors regulate plant secondary metabolites. Plants, 12(3), 447. https://doi.org/10.3390/plants12030447
Rizvi, A., Ahmed, B., Khan, M. S., El-Beltagi, H. S., Umar, S., & Lee, J. (2022). Bioprospecting plant growth promoting rhizobacteria for enhancing the biological properties and phytochemical composition of medicinally important crops. Molecules, 27(4), 1407. https://doi.org/10.3390/molecules27041407
Salehi, B., Zakaria, Z. A., Gyawali, R., Ibrahim, S. A., Rajkovic, J., Shinwari, Z. K., Khan, T., Sharifi-Rad, J., Ozleyen, A., Turkdonmez, E., Valussi, M., Tumer, T. B., Monzote Fidalgo, L., Martorell, M., & Setzer, W. N. (2019). Piper species: A comprehensive review on their phytochemistry, biological activities and applications. Molecules, 24(7), 1364. https://doi.org/10.3390/molecules24071364
Sander, A., Bival Štefan, M., Sander, T., Kučić Grgić, D., Parlov Vuković, J., Blažević, I., & Jablan, J. (2025). Characterization of essential oils and ethanolic extracts from nine pepper species: Antioxidant and antimicrobial activity and spectroscopic analysis. Molecules, 30(20), 4140. https://doi.org/10.3390/molecules30204140
Selwal, N., Rahayu, F., Herwati, A., Latifah, E., Supriyono, Suhara, C., Kade Suastika, I. B., Mahayu, W. M., & Wani, A. K. (2023). Enhancing secondary metabolite production in plants: Exploring traditional and modern strategies. Journal of Agriculture and Food Research, 14, 100702. https://doi.org/10.1016/j.jafr.2023.100702
Shabrina, A. M., Azzahra, R. S. S., Permata, I. N., Dewi, H. P., Safitri, R. A., Maya, I., Aulia, R. N., Sriwidodo, S., Mita, S. R., Amalia, E., & Putriana, N. A. (2025). Potential of natural-based sun protection factor (SPF): A systematic review of curcumin as sunscreen. Cosmetics, 12(1), 10. https://doi.org/10.3390/cosmetics12010010
Shah, S. S., Shi, L., Li, Z., Ren, G., Zhou, B., & Qin, P. (2020). Yield, agronomic and forage quality traits of different quinoa (Chenopodium quinoa Willd.) genotypes in Northeast China. Agronomy, 10(12), 1908. https://doi.org/10.3390/agronomy10121908
Shahzad, M., Hayat, R., Mujtaba, G., Rehman, W. U., & Nadeem, M. (2025). Biofertilizers in sustainable agriculture: Mechanisms, applications, and future prospects. Discover Agriculture, 3(1), 224. https://doi.org/10.1007/s44279-025-00318-0
Sholehah, D. N., & Ulya, M. (2023). Javanese long pepper (Piper retrofractum Vahl.): Botanical, cultivation, post-harvest and utilization review. Journal of Science in Agrotechnology, 1(2), 76–88. https://doi.org/10.21107/jsa.v1i2.15
Stojanović-Radić, Z., Pejčić, M., Dimitrijević, M., Aleksić, A., Kumar, N. V. A., Salehi, B., Cho, W. C., & Sharifi-Rad, J. (2019). Piperine—A major principle of black pepper: A review of its bioactivity and studies. Applied Sciences, 9(20), 4270. https://doi.org/10.3390/app9204270
Takshak, S., & Agrawal, S. B. (2019). Defense potential of secondary metabolites in medicinal plants under UV-B stress. Journal of Photochemistry and Photobiology B: Biology, 193, 51–88. https://doi.org/10.1016/j.jphotobiol.2019.02.002
Tang, Y., Nian, L., Zhao, X., Li, J., Wang, Z., & Dong, L. (2025). Bio-organic fertilizer application enhances silage maize yield by regulating soil physicochemical and microbial properties. Microorganisms, 13(5), 959. https://doi.org/10.3390/microorganisms13050959
Timofeeva, A. M., Galyamova, M. R., & Sedykh, S. E. (2023). Plant growth-promoting soil bacteria: Nitrogen fixation, phosphate solubilization, siderophore production, and other biological activities. Plants, 12(24), 4074. https://doi.org/10.3390/plants12244074
Venkatasai, N. N., Shetty, D. N., Vinay, C. M., Sekar, M., Muthusamy, A., & Rai, P. S. (2025). A comprehensive review of factors affecting growth and secondary metabolites in hydroponically grown medicinal plants. Planta, 261(3), 48. https://doi.org/10.1007/s00425-025-04619-y
Wadood, S. A., Jiang, Y., Nie, J., Li, C., Rogers, K. M., Liu, H., Zhang, Y., Zhang, W., & Yuan, Y. (2023). Effects of light shading, fertilization, and cultivar type on the stable isotope distribution of hybrid rice. Foods, 12(9), 1832. https://doi.org/10.3390/foods12091832
Wan, Y., Zhou, M., Le, L., Gong, X., Jiang, L., Huang, J., Cao, X., Shi, Z., Tan, M., Cao, Y., Wu, X., Wu, Q., Ye, X., Liu, C., Li, X., Peng, L., Xiang, D., & Zou, L. (2022). Evaluation of morphology, nutrients, phytochemistry and pigments suggests the optimum harvest date for high-quality quinoa leafy vegetable. Scientia Horticulturae, 304, 111240. https://doi.org/10.1016/j.scienta.2022.111240
Wu, W., Chen, L., Liang, R., Huang, S., Li, X., Huang, B., Luo, H., Zhang, M., Wang, X., & Zhu, H. (2025). The role of light in regulating plant growth, development and sugar metabolism: A review. Frontiers in Plant Science, 15, 1507628. https://doi.org/10.3389/fpls.2024.1507628
Xing, Y., Xie, Y., & Wang, X. (2025). Enhancing soil health through balanced fertilization: A pathway to sustainable agriculture and food security. Frontiers in Microbiology, 16, 1536524. https://doi.org/10.3389/fmicb.2025.1536524
Yu, J.-B., Bai, M., Wang, C., Wu, H., & Liang, X. (2024). Regulation of secondary metabolites accumulation in medicinal plants by rhizospheric and endophytic microorganisms. Medicinal Plant Biology, 3(1), e011. https://doi.org/10.48130/mpb-0024-0011
Zhang, H., Zhao, Q., Wang, Z., Wang, L., Li, X., Fan, Z., Zhang, Y., Li, J., Gao, X., Shi, J., & Chen, F. (2021). Effects of nitrogen fertilizer on photosynthetic characteristics, biomass, and yield of wheat under different shading conditions. Agronomy, 11(10), 1989. https://doi.org/10.3390/agronomy11101989
Zhang, S., Zhang, L., Zou, H., Qiu, L., Zheng, Y., Yang, D., & Wang, Y. (2021). Effects of light on secondary metabolite biosynthesis in medicinal plants. Frontiers in Plant Science, 12, 781236. https://doi.org/10.3389/fpls.2021.781236
Zhao, Y., Liu, G., Yang, F., Liang, Y., Gao, Q., Xiang, C., Li, X., Yang, R., Zhang, G., Jiang, H., Yu, L., & Yang, S. (2023). Multilayered regulation of secondary metabolism in medicinal plants. Molecular Horticulture, 3(1), 11. https://doi.org/10.1186/s43897-023-00059-y
Ziegenhagen, R., Heimberg, K., Lampen, A., & Hirsch-Ernst, K. I. (2021). Safety aspects of the use of isolated piperine ingested as a bolus. Foods, 10(9), 2121. https://doi.org/10.3390/foods10092121
Zou, R., Zhou, Y., Lu, Y., Zhao, Y., Zhang, N., Liu, J., Zhang, Y., & Fu, Y. (2024). Preparation, pungency and bioactivity transduction of piperine from black pepper (Piper nigrum L.): A comprehensive review. Food Chemistry, 456, 139980. https://doi.org/10.1016/j.foodchem.2024.139980
Copyright (c) 2026 Pramono Hadi, Riski Rahma Noviana, Suwardi, Agung Mugi Widodo
This work is licensed under a Creative Commons Attribution 4.0 International License.
Authors who publish with ISEJ: Indonesian Science Education Journal agree to the following terms:
- Authors retain copyright and grant the ISEJ: Indonesian Science Education Journal right of first publication with the work simultaneously licensed under Creative Commons Attribution License (CC BY 4.0) that allows others to share the work with an acknowledgment of the work's authorship and initial publication in this journal.
- Authors can enter into separate, additional contractual arrangements for the non-exclusive distribution of the published version of the work (e.g., post it to an institutional repository or edit it in a book), with an acknowledgment of its initial publication in this journal.
- Authors are permitted and encouraged to post their work online (e.g., in institutional repositories or on their website) before and during the submission process, as it can lead to productive exchanges, as well as earlier and greater citation of published work.
_(3).png)
