POLLINATION AND REPRODUCTIVE COMPATIBILITY TESTS IN CACAO (THEOBROMA CACAO L.) IN THE TRANSAMAZONIAN REGION

Authors

DOI:

https://doi.org/10.66104/1b5x5b09

Keywords:

controlled pollination; self-incompatibility; genetic compatibility; fruit set

Abstract

Cacao cultivation (Theobroma cacao L.) has high socioeconomic, environmental, and productive importance for tropical regions, especially within family farming systems. However, crop productivity is strongly limited by the low efficiency of natural pollination and the occurrence of genetic self-incompatibility mechanisms, which significantly reduce fruit set. In this context, the present systematic review aimed to describe and discuss the main aspects related to floral biology, pollination, and genetic compatibility in cacao, with emphasis on the artificial pollination methodology used to determine sexual compatibility among genotypes. The analyzed literature demonstrates that cacao possesses highly specialized flowers, mainly dependent on insects of the family Ceratopogonidae for natural pollination. Nevertheless, environmental factors, ecological limitations, and late-acting self-incompatibility mechanisms compromise the reproductive efficiency of the species, resulting in low fruiting rates. The review also demonstrates that artificial pollination constitutes an essential tool for studies on genetic compatibility, selection of superior genotypes, and breeding programs. The procedure involves standardized stages of flower selection and isolation, pollen collection, manual pollination, and post-pollination monitoring, allowing the evaluation of fruit set under controlled conditions. Furthermore, self-compatible genotypes exhibit greater productive stability and lower dependence on natural pollinators, making them strategic for commercial plantations. It is concluded that knowledge regarding reproductive biology and genetic compatibility is essential for optimizing breeding programs, increasing reproductive efficiency, and contributing to the productive sustainability of cacao cultivation. Additionally, the reviewed studies indicate that understanding intercompatibility relationships among clones allows more efficient planning of polyclonal arrangements, favoring cross-fertilization, fruit formation, and productive stability in orchards. Thus, the standardization of artificial pollination protocols represents technical support for reproductive research and crop management strategies. These methodologies also contribute to the identification of compatible genetic materials with agronomic potential, supporting orchard establishment, reducing reproductive failures, and improving pollination efficiency under tropical environmental conditions characterized by climatic and variability.

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Author Biographies

  • Sidevaldo Santana de Jesus, Federal Rural University of the Amazon (UFRA)

    Agricultural Technician from the Regional Agricultural High School of CEPLAC (EMARC). Holds a degree in Pedagogy from the State University of Pará (UEPA). Master's degree in Agronomy from the Federal Rural University of the Amazon (UFRA). Currently works as a Rural Extension Agent II at the Technical Assistance and Rural Extension Company of the State of Pará (EMATER-PA) and the Xingu Sustainable Regional Development Plan. Has experience in the field of Agronomy, with an emphasis on Rural Extension.

  • Harleson Sidney Almeida Monteiro, Sao Paulo State University (UNESP), Brazil

    Agronomist graduated from the Federal Rural University of the Amazon (UFRA). PhD candidate and Master in Agronomy (Horticulture) from the São Paulo State University "Júlio de Mesquita Filho". He also holds a degree in Management Processes from Paulista University (UNIP), a Technical degree in Occupational Safety from the Federal Institute of Education, Science and Technology of Pará (IFPA), and a Technical degree in Forestry from the Juscelino Kubitschek de Oliveira Agroindustrial School. During his studies, he worked as a scholarship monitor for the Genetics discipline between 2019 and 2020, and as a volunteer monitor for the disciplines of Fruit Growing, Vegetable Growing and Tropical Fruit Growing. He is a member of the Research Group on Fruit Production in the Amazon - FRUTAM, focusing on the management and improvement of native fruit trees in the region. He participated as a scholarship recipient in the PIBIC/CNPq program and was a student in the National Virtual Mobility Program Promover ANDIFES in the Agronomy course in the first semester of 2021. He contributes as a collaborator to the Agricultural Residency of MAPA/UFRA, where he works with Integrated Technologies in Family Farming, promoting the dissemination of knowledge and sustainable development in the Amazon. He has experience in rural extension, family farming, and organic fruit production, specializing in Horticulture and Fruit Growing, especially in management and cultural practices in fruit trees. His main areas of interest include native Amazonian fruit trees (such as açaí, bacabi, bacuri, cupuaçu, pupunha), tropical fruit trees (such as papaya, banana, soursop, acerola, guava, rambutan), and temperate and subtropical fruit trees (such as grapevine, avocado, persimmon, peach), focusing on pruning, rootstocks, training systems, and plant growth regulators.

  • Larissa Pacheco Nogueira, Federal Rural University of the Amazon (UFRA)

    Agricultural Engineer, graduated from the Federal Rural University of the Amazon (UFRA). Actively participates in the FRUTAM research group, focused on the study and development of technologies applied to Amazonian fruit trees, with an emphasis on intercropping systems and sustainable management. Has experience in environmental regularization, acquired during an internship at the Flor da Mata consultancy, working on the monitoring of Rural Activity License (LAR) and Rural Environmental Registry (CAR) processes. Throughout this period, she participated in several extension projects and technical events, such as Expofruta on the Belém and Igarapé-Açu campuses, and Show Agro in Paragominas, contributing to the integration between the university, producers, and the local community. Her interests lie in the areas of Amazonian agroecosystems, tropical fruit growing, sustainable management, and technological innovation in rural areas.

  • Sinara de Nazaré Santana Brito, Sao Paulo State University (UNESP)

    Agricultural Engineer from the Federal Rural University of the Amazon (UFRA), Master in Agronomy (Horticulture) from the São Paulo State University – Faculty of Agronomic Sciences (UNESP/FCA) and currently a doctoral candidate at the same institution, affiliated with the Department of Plant Production (Horticulture). She holds a technical degree in Occupational Safety from the Federal Institute of Pará (IFPA). She works in the field of Horticulture, researching post-harvest biochemistry and physiology of fruit trees, focusing on quality conservation, metabolism of bioactive compounds, and the application of plant growth regulators. She collaborated with the MAPA/UFRA Agricultural Residency program, in the area of ​​Integrated Technologies in Family Farming, with work focused on disseminating sustainable practices in the Amazon. She is a member of the Research Group on Fruit Growing in the Amazon (FRUTAM), the Technological Innovations in the Development of Plant-Based Products group (GITPV), the Interdisciplinary Research Group on Agroecosystems for Rural Development in the Tocantins Amazon (PIADRAT), and the Research Group on Fruit Growing (Frut!). Her work focuses on the interface between agricultural sustainability and food security, integrating approaches in plant metabolism and post-harvest physiology to understand the biochemical processes that determine the quality, conservation, and value of products derived from biodiversity.

  • Viviandra Manuelle Monteiro de Castro Trindade, Sao Paulo State University (UNESP)

    Agricultural Engineer from the Federal Rural University of the Amazon (UFRA). Master's degree in Environmental Sciences from the State University of Pará (UEPA), with research on the processing and consumption of açaí and its socioeconomic relevance in the Amazon. PhD candidate in Agronomy (Horticulture) at UNESP, working in the area of ​​horticultural species production and researching the fig production chain. Occupational Safety Technician from the Magalhães Barata Technical School (ETEMB). Former PIBIC/CNPq scholarship holder at the Museu Paraense Emílio Goeldi, working in myrmecology applied to fruit growing. Member of the research groups Fruit Growing in the Amazon (FRUTAM) and Teaching, Research and Extension Group in Fruit Growing (FRUTI/UNESP).

  • Cinthia Mota Veiga, Federal Rural University of the Amazon (UFRA)

    Agricultural Engineer from the Federal Rural University of the Amazon (UFRA). Works in the area of ​​Amazonian fruit farming, with expertise especially in cocoa and açaí crops. Participated in scientific initiation projects and activities focused on sustainable agriculture in the Amazon.

  • Nayara Ferreira Barros da Silva, Flor da Mata Environmental Services

    Agricultural Engineer (UFRA). Specialist in Integrated Pest Management, Georeferencing of Rural Properties, and MBA in Agribusiness Management. Environmental coordinator at Flor da Mata Environmental Services, working directly in the environmental regularization of rural properties, monitoring processes for the Rural Environmental Registry (CAR), Environmental Regularization Program (PRA), Recovery Plan for Degraded and/or Altered Areas, and Rural Activity License (LAR). She was a scholarship recipient in the Professional Agricultural Residency program (UFRA/MAPA). Member of the FRUTAM research group.

  • Deivid Almeida de Jesus, Federal University of Rio de Janeiro (UFRJ)

    Bachelor of Science in Biological Sciences from the Federal University of Western Pará (UFOPA). Master of Science in Biological Sciences (Genetics) and PhD candidate in the same program at the Federal University of Rio de Janeiro (UFRJ). Has an interest and experience in plant genetics and bioinformatics.

  • Glenda Rafaele da Silva, Federal Rural University of the Amazon (UFRA)

    Undergraduate student in Agronomy at the Federal Rural University of the Amazon (UFRA). Member of FRUTAM – Research Group on Fruit Growing in the Amazon, focusing on native and exotic fruit growing in the Amazon.

  • Antonia Benedita da Silva Bronze, Federal Rural University of the Amazon (UFRA)

    She was Pro-Rector of Extension (PROEX/UFRA) at the Federal Rural University of the Amazon (UFRA). She holds a degree in Agronomy from the Federal Rural University of the Amazon (UFRA); she was a scientific initiation scholarship holder from 1999 to 2002 at the same institution; a Master's degree in Agronomy, in the area of ​​concentration Tropical Plant Biology, from the Federal Rural University of the Amazon (UFRA); a Doctorate in Agricultural Sciences, in the area of ​​concentration Amazonian agroecosystems, from the Federal Rural University of the Amazon (UFRA); she was a substitute professor at the Federal Rural University of the Amazon, teaching the Genetics discipline from 2005 to 2007; she was a rural extension agent I - agronomist at the Technical Assistance and Rural Extension Company of Pará - EMATER-PA, from 2006 to 2010; and she was the coordinator of the local EMATER-PA office in the municipality of São João de Pirabas. Coordinator of a research project for a tropical fruit growing hub in the State of Pará: Implementation of a network of solidarity enterprises for family farmers in the Salgado Microregion - CNPQ/MDA; In 2010, she was appointed to the staff of the Federal Rural University of the Amazon as a professor at the 3rd level, with exclusive dedication in the area of ​​Fruit Growing, Seed and Seedling Technology, and Horticulture, approved through a public competition. She was the coordinator of the postgraduate specialization in Low Carbon Agriculture in the Amazon, through the Amapá Project. Member of the Açaí Route Committee, BR-316 Hub, through the National Integration Routes Program of the Ministry of Regional Development (MDR). She was the coordinator of the Agricultural Residency in the project INTEGRATED TECHNOLOGIES IN FAMILY AGRICULTURE: Dissemination of knowledge, expertise and sustainable development in the Amazon (2020 to 2023). Member of the State Council for Sustainable Rural Development of Pará. Tenured Professor and Researcher in the Agronomy Graduate Program (PGAGRO) at UFRA, in the area of ​​phytotechnics, teaching the subject of fruit production systems in the Amazon and conducting research on Amazonian fruits and sustainability. Coordinator of the research and extension group in fruit growing in the Amazon, FRUTAM, registered with CNPQ. Her main areas of expertise include: Fruit growing, Amazonian fruit growing, açaí cultivation, cocoa cultivation, agroforestry systems, plant genetics, genetic resources, native Amazonian palm trees, vegetative propagation, and production and marketing systems for Amazonian fruits.

References

AMARAL, C. A. do; FREITAS NORONHA, R. H. de; PASSOS, M. M. S.; CAMPOS, D. O.; LIMA SILVA, N. Produtividade do cacau no sul da Bahia. Revista de Política Agrícola, Brasília, v. 33, e01957, 2025.

BAHIA, R. C.; CORRÊA, R. X.; SANTOS, R. C.; MACHADO, R. C. R.; LUZ, E. D. N.; ARAÚJO, I. S.; AHNERT, D. Inheritance of the number of ovules per ovary and selection of cacao genotypes. American Journal of Plant Sciences, v. 4, p. 1387–1392, 2013. DOI: http://dx.doi.org/10.4236/ajps.2013.47169 DOI: https://doi.org/10.4236/ajps.2013.47169

BARROSO, D. F. R.; BERRÊDO, J. F.; PIMENTEL, M. A. S. da; MOURA, M. do N.; ALMEIDA, R. P. S.; THALÊS, M. C.; VILHENA, M. do P. S. P. Efeitos de variáveis hidroclimáticas sobre a produção de cacau (Theobroma cacao L.) em Mocajuba, Amazônia Oriental. Revista Brasileira De Geografia Física, v. 18, n. 05, p. 3558–3572, 2025. DOI: https://doi.org/10.26848/rbgf.v18.05.p3558-3572 DOI: https://doi.org/10.26848/rbgf.v18.05.p3558-3572

BEKELE, F.; PHILLIPS-MORA, W. Melhoramento genético do cacau (Theobroma cacao L.). In: AL-KHAYRI, J. M.; JAIN, S. M.; JOHNSON, D. V. (eds.). Advances in Plant Breeding Strategies: Industrial and Food Crops. Cham: Springer, 2019. p. 581–616. DOI: https://doi.org/10.1007/978-3-030-23265-8_12. DOI: https://doi.org/10.1007/978-3-030-23265-8_12

BILLES, D. J. Pollination of Theobroma cacao L. in Trinidad, B.W.I. Tropical Agriculture, v. 18, p. 151–156, 1941.

BRASIL. Ministério da Agricultura e Pecuária. Cacau do Brasil. Brasília: MAPA, 2020.

BRIDGEMOHAN, P.; MOHAMMED, M. The ecophysiology of abiotic and biotic stress on the pollination and fertilization of cacao (Theobroma cacao L.). In: OLIVEIRA, A. B. (ed.). Abiotic and Biotic Stress in Plants. London: IntechOpen, 2019. DOI: https://doi.org/10.5772/intechopen.84528

CARLETTO, G. A.; SORIA V., J. Testes de graus de autocompatibilidade em cacaueiro (Theobroma cacao L.). Revista Theobroma, v. 3, p. 26–35, 1973.

CEPLAC – COMISSÃO EXECUTIVA DO PLANO DA LAVOURA CACAUEIRA. Cacau: botânica, morfologia, fisiologia, produção e processamento. Ilhéus: CEPLAC, 2020.

CORONADO-ALEANS, V.; ARGUELLO CARREÑO, E. F.; MORALES GOMEZ, J. E. Sexual compatibility assessment of two cocoa genotypes (Theobroma cacao L.) in Santander, Colombia. Revista de Ciencias Agrícolas, Pasto, v. 42, n. 2, e2269, 2025. DOI: https://doi.org/10.22267/rcia.20254202.269. DOI: https://doi.org/10.22267/rcia.20254202.269

DANI, D.; ROKHMAH, D. N. A review of the role of pollination on the yield of cocoa plant. Jurnal Kultivasi, v. 21, n. 3, p. 249–260, 2022. DOI: https://doi.org/10.24198/kultivasi.v21i3.41513. DOI: https://doi.org/10.24198/kultivasi.v21i3.41513

FRIMPONG-ANIN, K.; ADJALOO, M. K.; KWAPONG, P. K.; ODURO, W. Structure and stability of cocoa flowers and their response to pollination. Journal of Botany, v. 2014, Article ID 513623, 2014. DOI: https://doi.org/10.1155/2014/513623. DOI: https://doi.org/10.1155/2014/513623

GALVÃO, E. L.; SILVESTRE, J. V.; CRUZ, J. N.; SILVA, N. C. S. da. Cacau (Theobroma cacao): uma visão geral de pesquisas atuais sobre aspectos botânicos, fitoquímicos e farmacológicos. Research, Society and Development, v. 13, n. 5, e8113545810, 2024. DOI: http://dx.doi.org/10.33448/rsd-v13i5.45810.

GODOY, P. R. E.; SOUZA, M. M.; ROZA, F. A.; LAWINSCKY, P. R.; ARAÚJO, I. S.; AHNERT, D. Performance polínica em cacaueiros (Theobroma cacao L.) autocompatíveis e autoincompatíveis. Brazilian Journal of Botany, São Paulo, v. 32, n. 3, p. 617–620, 2009. DOI: https://doi.org/10.1590/S0100-84042009000300019 DOI: https://doi.org/10.1590/S0100-84042009000300019

JARAMILLO, M. A.; REYES-PALENCIA, J.; JIMÉNEZ, P. Floral biology and flower visitors of cocoa (Theobroma cacao L.) in the upper Magdalena Valley, Colombia. Flora, v. 313, 152480, 2024. DOI: https://doi.org/10.1016/j.flora.2024.152480. DOI: https://doi.org/10.1016/j.flora.2024.152480

LANAUD, C.; FOUET, O.; LEGAVRE, T.; LOPES, U.; SOUNIGO, O.; EYANGO, M. C.; CLÉMENT, D. Deciphering the Theobroma cacao self-incompatibility system: from genomics to diagnostic markers for self-compatibility. Journal of Experimental Botany, Oxford, v. 68, n. 17, p. 4775–4790, 2017. DOI: https://doi.org/10.1093/jxb/erx293. DOI: https://doi.org/10.1093/jxb/erx293

LÓPEZ, M. E.; RAMIREZ, O. A.; DUBON, A.; RIBEIRO, T. H. C.; DIAZ, F. J.; CHALFUN-JUNIOR, A. Sexual compatibility in cacao clones drives arrangements in the field leading to high yield. Scientia Horticulturae, Amsterdam, v. 287, 110276, 2021. DOI: https://doi.org/10.1016/j.scienta.2021.110276. DOI: https://doi.org/10.1016/j.scienta.2021.110276

RÍOS-MOYANO, D. K.; RODRÍGUEZ-CRUZ, F. A.; HORMAZA-MARTÍNEZ, P. A.; RAMÍREZ-GODOY, A. Characterization of pollinators associated with cocoa cultivation and their relationship with natural effective pollination. Diversity, Basel, v. 17, n. 3, p. 189, 2025. DOI: https://doi.org/10.3390/d17030189. DOI: https://doi.org/10.3390/d17030189

SOUZA, M. S.; VENTURIERI, G. A. Floral biology of cacauhy (Theobroma speciosum – Malvaceae). Brazilian Archives of Biology and Technology, Curitiba, v. 53, n. 4, p. 861–872, 2010. DOI: https://doi.org/10.1590/S1516-89132010000400016 DOI: https://doi.org/10.1590/S1516-89132010000400016

VANSYNGHEL, J.; THOMAS, E.; OCAMPO-ARIZA, C.; MAAS, B.; ULLOQUE-SAMATELO, C.; ZHANG, D.; STEFFAN-DEWENTER, I. Cross-pollination with native genotypes improves fruit set and yield quality of Peruvian cacao. Agriculture, Ecosystems and Environment, Amsterdam, v. 357, 108671, 2023. DOI: https://doi.org/10.1016/j.agee.2023.108671 DOI: https://doi.org/10.1016/j.agee.2023.108671

YAMADA, M. M.; FALEIRO, F. G.; SANTOS, R. F. dos; PIRES, J.; YAMADA, M. M.; DOS SANTOS, R. F.; PIRES, J. L. Relação entre incompatibilidade, cor de frutos, incidência de vassoura-de-bruxa e a produção de frutos em cacaueiro. Agrotropica, Ilhéus, v. 26, n. 3, p. 207–210, 2014. DOI: https://doi.org/10.21757/0103-3816.2014v26n3p207-210

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Published

2026-05-18

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Vol. 13 No. 10 (2026): REMUNOM - ISSN 2178-6925

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Artigos

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Copyright (c) 2026 Sidevaldo Santana de Jesus, Harleson Sidney Almeida Monteiro, Larissa Pacheco Nogueira, Sinara de Nazaré Santana Brito, Viviandra Manuelle Monteiro de Castro Trindade, Cinthia Mota Veiga, Nayara Ferreira Barros da Silva, Deivid Almeida de Jesus, Glenda Rafaele da Silva, Antonia Benedita da Silva Bronze, Gleicilene Brasil de Almeida

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How to Cite

POLLINATION AND REPRODUCTIVE COMPATIBILITY TESTS IN CACAO (THEOBROMA CACAO L.) IN THE TRANSAMAZONIAN REGION. (2026). REMUNOM, 13(10), 1-25. https://doi.org/10.66104/1b5x5b09