Genotype and within-pod bean position microenvironment effect on seed choice for raising cocoa (Theobroma cacao L.) seedlings

Daniel B. ADEWALE, Beatrice A. NDUKA


The probable role of within-pod microenvironment on seed sizes, seedling vigour and biomass yield of four cocoa genotypes was investigated for two years. The respective main, sub and sub-sub plots in the split-split plot experimental design were years, genotypes and within-pod bean positions. Data were taken on cocoa bean length, width and thickness after each pod was opened. Four weekly periodic data were obtained for plant height (PH), stem girth (SG) and number of leaves (NOL); root and shoot biomass yield were also recorded. Analysis of variance revealed significant (p ≤ 0.05) bean position, genotypes, years and some interaction on the studied traits. Means of the levels of the three factors differed significantly (p ≤ 0.05). Proximal, middle and distal positions were distinct within-pod microenvironments. The pod middle cavity housed the longest, widest and heaviest beans. Trend analysis of the growing sequences of NOL, PH and SG by the four genotypes differed with bean locations. For bean length, GGE biplot respectively identified CRIN Tc1, CRIN Tc2 and CRIN Tc3 as the best genotype for middle, proximal and the distal positions. The intra-locular space within the pod enhanced differential seed development and maturation; this was evident in the seedling vigour.


bean position; cocoa; micro-environments; GGE biplot; seedling vigour

Full Text:



Adewale, B.D., Okonji, C., Oyekanmi, A.A., Akintobi, D.A.C. and Aremu, C.O. (2010). Genotypic variability and stability of some grain yield components of cowpea. African Journal of Agricultural Research, 5, 874-880.

Adewale, B.D., Adeigbe, O.O. and Muyiwa, A.A. (2016). Cocoa seed garden: a means to disseminating improved planting materials for enhanced national productivity: A review. Agricultural Reviews, 37, 205-212.

Amma, S.P., Mininol, J.S. and Bai, L.E.S. (2011). Utilization of introduced genetic resources in cocoa. Cashew Cocoa Journal, 3, 16 -18.

Baiyeri, K.P. (2006). Seedling emergence and growth of pawpaw (Carica papaya) grown under different coloured shade polyethylene. International Journal of Agrophysics, 20, 77-84.

Bekele, F.L., Bekele, I., Butler, D.R. and Bidaisee, G.G. (2006). Patterns of morphological variation in a sample of cacao (Theobroma cacao L.) from the International Cocoa Genebank, Trinidad. Genetic Resources and Crop Evolution, 53, 933–948.

Bennett, E.J., Roberts, J.A. and Wagstaff, C. (2011). The role of the pod in seed development: strategies for manipulating yield. New Physiologist, 190, 838–853.

Chacon, P., Ramiro, B. and Carolina, H. (1998). The Effect of seed size on germination and seedling growth of Cryptocarya alba (Lauraceae) in Chile. Revista Chiliena de Historia Natural, 71, 189-197.

Clark, M.D. and Watkins, E. (2012). Broad-sense heritability estimates of turfgrass performance characteristics in native Prairie june grass germplasm. Horticultural Sciences, 47, 1228-1233.

Copeland, L.O. and McDonald, M.B. (2001). Seed vigour and vigour tests. In L.O. Copeland & M.B. McDonald (Eds.), Principles of Seed Science and Technology (pp. 121-144). New York: Kluwer Academic Publishing Group.

CRIN (2011). New hybrid cocoa varieties for Nigeria: Attribute and field management requirements. Cocoa Research Institute of Nigeria, Idi-Ayunre, Ibadan. Library Information and Documentation Department.

Crossa, J., Fox, P.N., Pfeiffer, W.H., Rajaram, S. and Gauch, H.G. (1991).

AMMI adjustment for statistical analysis of an international wheat yield trial. Theoretical and Applied Genetics, 81, 27-37.

Cruz-Garcia, F., Gonzalez-Hernandez, V.A., Molina-Moreno, J. and Vazquez-Ramos, J.M. (1995). Seed deterioration and respiration as related to DNA metabolism in germinating maize. Seed Science and Technology, 23, 477-486.

Enayatgholizadeh, M.R., Alami-Saeid, K.H., Bakhshandeh, A.M., Dehghan-Shoar, M., Ghaineh, M.H. and Sharafizadeh, M. (2011). Response of the morphologic characteristics of S.C704 maize affected by the source and seed size in Khuzestan. Australian Journal of Basic and Applied Sciences, 5, 369-374.

Ghassemi-Golezani, K., Bakhshy, J., Raey, Y. and Hossinzadeh-Mahootchy, A. (2010). Seed vigour and field performance of winter oilseed rape (Brassica napus L.) cultivars. Notulae Botanicae Horti Agrobotanici Cluj-Napoca, 38, 146-50.

Giles, B.E. (1990). The effects of variation in seed size on growth and reproduction in the wild barley Hordeum vulgare ssp. spontaneum. Heredity, 64, 239-250.

Hammed, L.A., Olaiya, A.O., Lawal, I.O., Idowu, O.T.H. and Aiyelaagbe, I.O.O. (2013). Effects of some bean characters on germination and seedling growth of cocoa (Theobroma cacao L.). Nigerian Journal of Horticultural Sciences, 17,126-134.

Ibikunle, B.O. (1967). Effect of position of beans in the pod on the performance of cocoa seedlings. Cocoa Research Institute of Nigeria Annual Report.

Illipronti Jr., R.A., Lommen, W.J.M., Langerak, C.J. and Struik, P.C. (2000). Time of pod set and seed position on the plant contribute to variation in quality of seeds within soybean seed lots. Netherlands Journal of Agricultural Sciences, 48, 165-180.

Iremiren, G.O., Famaye, A.O. and Oloyede, A.A. (2007). Effects of pod sizes and bean positions in pod on the germination and seedling growth of cocoa (Theobroma cacao). African Crop Science Conference Proceedings, 8, 1979-1982.

Kaushik, N., Kumar, K., Kumar. S., Kaushik, N. and Roy, S. (2007). Genetic variability and divergence studies in seed traits and oil content of Jatropha (Jatropha curcas) accessions. Biomass and Bioenergy, 31, 497-502.

Khan, D., Sahito, Z.A., Zaki, M.J. and Shaukat, S.S. (2014). Axial dimensions of pods and seeds and within-pod-allocation of phytomass and seed packaging cost in Erythrina suberosa Roxb. (Papilionaceae). International Journal of Biology and Biotechnology, 11, 191-206.

Lee, T.D. (1988). Patterns of fruit and seed production. In J. Lovett-Doust (Eds.), Plant reproductive ecology: patterns and strategies (pp 179–202). New York NY: Oxford University Press.

Matthews, S., Noli, E., Demir, I., Khajeh, H.M. and Wagner, M.H. (2012). Evaluation of seed quality: from physiology to international standardization. Seed Science Research, 22, 69-73.

Moreno-Martinez, E., Vazquez-Badillo, M.E., Rivera, A., Navarrete, R. and Fasquivel-Villagrana, F. (1998). Effect of seed shape and size on germination of corn (Zea mays L.) stored under adverse conditions. Seed Science and Technology, 26, 439-48.

Mortazavian, S.M.M. and Azizinia, S. (2014). Nonparametric stability analysis in multi-environment trial of canola. Turkish Journal Field Crops, 19, 108-117.

Motamayor, J.C., Lachenaud, P., Da Silva, E., Mota, J.W., Loor, R., Kuhn, D.N., Brown, J.S. and Schnell, R.J. (2008). Geographic and genetic population differentiation of the Amazonian chocolate tree (Theobroma cacao L.). PLOS ONE 3(10), e3311.

Nakamura, R.R. (1988). Seed abortion and seed size variation within fruits of Phaseolus vulgaris: pollen donor and resource limitation effects. American Journal of Botany, 75, 1003-1010.

Omokhafe, K.O. and Alika, J.E. (2004). Clonal variation and correlation of seed characters in Hevea brsiliensis (Muell.) Arg. Industrial Crops Production, 19, 175-184.

Opoku-Ameyaw, K., Baah, F., Gyedu-Akoto, E., Anchirinah, V., Dzahini-Obiately, H., Cudjoe, A.R., Aquaye, S. and Opoku, S.Y. (2010). Cocoa manual: A source book for sustainable cocoa production. New Tafo-Akim, Cocoa Research Institute of Ghana (CRIG).

Ortiz, R. and Ng, N.Q. (2000). Genotype × Environment interaction and its analysis germplasm characterization and evaluation. In I.J. Ekanayake and R. Ortiz, (Eds.), Genotype × Environment interaction analysis of IITA mandate crops in Sub-Saharan Africa (pp 32-40). Ibadan: IITA.

Ortiz, F. (2016). Theobroma cacao L. Monograph. Colegio Bolivar.

Pacheco, A., Vargas, M., Alvarado, G., Rodríguez, G., Crossa, J. and Burgueño, J. (2016). GEA-R (genotype environment analysis with R for Windows). Version 4.1. International Maize and Wheat Improvement Center.

Perin, A., Araújo, A.P. and Teixeira, M.G. (2002). Efeito do tamanho da semente na acumulação de biomassa e nutrientes e na produtividade do feijoeiro. Pesquisa Agropecuária Brasileira, 37, 1711-1718.

Phillips‐Mora, W., Arciniegas‐Leal, A., Mata‐Quiros, A. and Motamayor‐Arias, J.C. (2013). Catalogue of cacao clones selected by CATIE for commercial plantings, Turrialba.

R Development Core Team. (2013). R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Austria. URL Retrieved from

SAS. (2011). SAS Online Doc® 9.4. SAS Institute Inc., Cary, NC.

Singh, R.K. and Chaudhary, B.D. (1999). Biometrical methods in quantitative genetic analysis. New Delhi: Kalyani Publisher.

Singh, N. and Pokhriyal, T.C. (2001). Variations in pod and seed traits in six different Dalbergia sissoo seed sources. Journal of Tropical Forest Science, 13, 162-170.

Susko, D.J. and Lovett-Doust L. (2000). Patterns of seed mass variation and their effects on seedling traits in Alliaria petiolata (Brassicaceae). American Journal of Botany, 87, 56–66.

Yan, W. and Kang, M.S. (2003). GGE biplot analysis: a graphical tool for breeders, geneticists and agronomists. Boca Raton, Florida Fl: CRC Press.



  • There are currently no refbacks.

Copyright (c) 2020 Daniel B. Adewale

Creative Commons License
This work is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License.


Acta agriculturae Slovenica is an Open Access journal published under the terms of the Creative Commons CC BY License.


eISSN 1854-1941