Scarification and doses of Acadian®, Stimulate® and Trichoderma spp. promote dormancy overcoming in Hymenaea courbaril L. seeds?

Oscar José Smiderle Aline das Graças Souza About the authors

Abstract:

The present study establishes the following research problem: Can different concentrations of Acadian®, Stimulate®, and Trichoderma spp. be effective in promoting the emergence and vigor of Hymenaea courbaril seedlings with or without mechanical scarification? The experimental design used was completely randomized, in a 4 x 3 x 2 factorial scheme, corresponding to four concentrations (0, 5.0, 10.0, and 15.0 mL.L-1), three hormones (Acadian®, Stimulate® and Trichoderma spp.), with and without mechanical scarification, with four replicates, each of which consisting of 25 seeds. Acadian® at a concentration of 5.0 mL.L-1 in scarified seeds of H. courbaril leads to a greater increase in the percentage of emergence and percentage of normal seedlings. Stimulate® at concentrations of 10.0 and 15.0 mL.L-1 is efficient in obtaining more than 85% emergence seedlings with seeds subjected to mechanical scarification. Trichoderma spp. from 9.0 mL.L-1 does not favor gains in the percentage of normal seedlings of Hymenaea courbaril grown from seeds with and without mechanical scarification. The dose of maximum technical efficiency (DMTE) of the bioregulator Acadian® is 8.50 mL.L-1 and, in scarified seeds of H. courbaril, it increases the percentage of seedling emergence.

Index terms:
Ascophyllum nodosum; bioregulators; mechanical scarification

Resumo:

O presente estudo estabelece o seguinte problema de pesquisa: Sementes com e sem escarificação mecânica sob diferentes concentrações de Acadian®, Stimulate® e Trichoderma ssp. podem ser eficazes na promoção de emergência e vigor de plântulas de Hymenaea courbaril? O delineamento experimental utilizado foi o inteiramente casualizado, em esquema fatorial 4 x 3 x 2, sendo as concentrações (0; 5,0; 10,0; e 15,0 mL.L-1), três hormônios (Acadian®, Stimulate® e Trichoderma ssp.) e com e sem escarificação mecânica, com quatro repetições, sendo cada uma delas composta de 25 sementes. O Acadian® na concentração de 5,0 mL.L-1 em sementes escarificadas de H. courbaril, exibe maior incremento no percentual de emergência e bem como na porcentagem de plântulas normais. O Stimulate® nas concentrações de 10,0 e 15,0 mL.L-1 é eficiente na obtenção de valores acima de 85% de emergência de plântulas com sementes submetidas a escarificação mecânica. Trichoderma spp. a partir de 9,0 mL.L-1 não favorece ganhos no percentual de plântulas normais de H. courbaril em sementes com e sem escarificação mecânica. A dose de máxima eficiência técnica (DMTE) do biorregulador Acadian® é de 8,50 mL.L-1 em sementes escarificadas de jatobá aumenta o percentual de emergência de plântulas.

Termos para indexação:
Ascophyllum nodosum; biorreguladores; escarificação mecânica

INTRODUCTION

Hymenaea courbaril L. belongs to the Fabaceae family and Caesalpinioideae subfamily, is a native forest species, semi-deciduous, belonging to the group of late secondary or climax species, and is used in popular medicine, civil construction, composition of heterogenous plantations and afforestation of parks and gardens (Costa et al., 2018COSTA, N.C.R.; CARVALHO, F.J.; JULIATTI, F.C.; CASTRO, M.V.; CUNHA, W.V. Diversity of fungi in seeds of Hymenaea stignocarpa and Hymenaea courbaril before and after fungicide treatments. Bioscience Journal, v.34, n.4, p.868-874, 2018. https://doi.org/10.14393/BJ-v34n1a2018-41866
https://doi.org/10.14393/BJ-v34n1a2018-4...
). In addition to the common name already mentioned, depending on the region in Brazil it is called jataí, jataí amarelo, jataí peba, jataí vermelho, jitaí, farinheira, jataíba, burandã, imbiuva, H. courbaril miúdo and jatobá da caatinga (Paixão et al., 2019PAIXÃO, M.V.S.; VIEIRA, K.M.; FERREIRA, E.A.; MÔNICO, A.F.; CARVALHO, A.J.C. Germination and dormity in jatobá seeds. International Journal of Advanced Engineering Research and Science, v.6, n.6, p.454-457, 2019. http://journal-repository.com/index.php/ijaers/article/view/578
http://journal-repository.com/index.php/...
).

According to Ristau et al. (2018RISTAU, A.C.P.; CRUZ, M.S.F.V.; REGO, C.A.R.M.; BRAZ, H.; OLIVEIRA, S.S.; COSTA, P.B.; MALAVASI, M.M.; MALAVI, C.U.; TSUTSUMI, C.Y. Biometric characterisation and physiological quality of seeds of Hymenaea courbaril L. Journal of Experimental Agriculture International, v.26, n.3, p.1-9, 2018. https://doi.org/10.9734/JEAI/2018/43824
https://doi.org/10.9734/JEAI/2018/43824...
), the fruit is composed of seeds (25 to 40% of weight), pod (50 to 70%), and pulp (5 to 10%). The protein value of H. courbaril fruit flour is similar to that of cornmeal and higher than that of cassava flour (Menezes-Filho et al., 2019MENEZES-FILHO, A.C.P.; SILVA, M.A.; PEREIRA, A.V.; OLIVEIRA FILHO, J. G.; CASTRO, C.F.S. Parâmetros físico-químicos, tecnológicos, atividade antioxidante, conteúdo de fenólicos totais e carotenóides das farinhas dos frutos do jatobá-do-cerrado (Hymenaea stigonocarpa Mart. ex Hayne). Multi-Science Journal, v.2, n.1, p.93-100, 2019. doi: 10.33837/msj.v2i1.900
https://doi.org/10.33837/msj.v2i1.900...
). It is worth highlighting that 100 grams of the fruit provides 115 calories, 29.4 grams of glucose, and 33 milligrams of vitamin C (Ramos et al., 2018RAMOS, F.S.A.R.; SANTOS, T.C.; FERREIRA, T.H.B.; GOMES, M.C.S.; MUNHOZ, C.L. Aceitabilidade de biscoito tipo cookie enriquecidos com farinha de jatobá. Cadernos de Agroecologia, v.13, n.2, p.1-7, 2018. http://cadernos.aba-agroecologia.org.br/index.php/cadernos/article/view/2311/2157
http://cadernos.aba-agroecologia.org.br/...
; Silva et al., 2019SILVA, C.P.; MANOLIO, S.F.R.A.; SAMPAIO, G.R.; BARROS, M.C.S.; NASCIMENTO, T.P.; CAMERON, L.C.; LARRAZ, M.S.F.; GOMES, J.A.A. Identification and action of phenolic compounds of Jatobá-do-cerrado (Hymenaea stignocarpa Mart.) on α-amylase and α-glucosidase activities and flour effect on glycemic response and nutritional quality of breads. Food Research International, v.116, n.18, p.1076-1083, 2019. https://pubmed.ncbi.nlm.nih.gov/30716891/
https://pubmed.ncbi.nlm.nih.gov/30716891...
), and the vitamin C concentration in H. courbaril fruit flour is similar to that of orange and 9 times higher than that of açaí pulp.

In addition to the medicinal relevance, it also has forest and environmental importance due to its potential as a carbon-fixing and -storing plant (Costa et al., 2018COSTA, N.C.R.; CARVALHO, F.J.; JULIATTI, F.C.; CASTRO, M.V.; CUNHA, W.V. Diversity of fungi in seeds of Hymenaea stignocarpa and Hymenaea courbaril before and after fungicide treatments. Bioscience Journal, v.34, n.4, p.868-874, 2018. https://doi.org/10.14393/BJ-v34n1a2018-41866
https://doi.org/10.14393/BJ-v34n1a2018-4...
). However, H. courbaril remains with no protection and the trees are threatened by predatory logging in most of the Amazon.

Besides affecting the long-term supply of wood, the destruction of H. courbaril trees can also reduce the supply of its important products. In the state of Roraima, the H. courbaril tree blooms between March and May and its fruiting occurs between August and October (Shanly and Medina, 2005SHANLEY, P.; MEDINA G. Frutíferas e plantas úteis na vida Amazônica- Jatobá Hymenaea courbaril. Belém: CIFOR, Imazon, 2005. 300 p.).

H. courbaril seeds are ovoid, with dark red color, glabrous, with an average fresh seed mass of 3.53 grams (Almeida et al., 2011ALMEIDA, M.B.; SOUZA, W.C.O.; GOMES, E.C.S.; RAMALHO, F.C.V. Descrição morfológica do fruto e semente do jatobá (Hymenaea courbaril L). Revista Semiárido de Visu, v.1, n.2, p.107-115, 2011. https://periodicos.ifsertao-pe.edu.br/ojs2/index.php/semiaridodevisu/article/view/39
https://periodicos.ifsertao-pe.edu.br/oj...
). H. courbaril seeds, when ripe, exhibit dormancy due to impermeability to water of the seed coat, so treatment is necessary to overcome dormancy, and a mechanical scarification is a form that allows increments in the percentage of seedling emergence (Duarte et al., 2016DUARTE, M.M.; PAULA, S.R.P.; FERREIRA, F.R.L.; NOGUEIRA, A.C. Morphological characterization of fruit, seed and seedling and germination of Hymenaea courbaril L. (Fabaceae) (‘Jatobá’). Journal of Seed Science, v.38, n.3, p.204-211, 2016. https://doi.org/10.1590/2317-1545v38n3159734
https://doi.org/10.1590/2317-1545v38n315...
).

Within these perspectives, mechanical scarification methods and bioregulators function as activators of metabolism and reactivate physiological processes in the different phases of germination, so they have been used as pre-germination treatments to optimize the germination of dormant seeds (Junges et al., 2016JUNGES, E.; MUNIZ, M.F.; MEZZOMO, R.; BASTOS, B.; MACHADO, R.T. Trichoderma spp. na produção de mudas de espécies florestais.Floresta e Ambiente , v.23, n.2, p.237-244, 2016. https://dx.doi.org/10.1590/2179-8087.107614
https://dx.doi.org/10.1590/2179-8087.107...
; Costa et al., 2018COSTA, N.C.R.; CARVALHO, F.J.; JULIATTI, F.C.; CASTRO, M.V.; CUNHA, W.V. Diversity of fungi in seeds of Hymenaea stignocarpa and Hymenaea courbaril before and after fungicide treatments. Bioscience Journal, v.34, n.4, p.868-874, 2018. https://doi.org/10.14393/BJ-v34n1a2018-41866
https://doi.org/10.14393/BJ-v34n1a2018-4...
).

Acadian® based on pure extracts of Ascophyllum nodosum seaweed stimulates radicle emergence, elongation, and growth of the piliferous region of the root system, resulting in better seedling establishment and water/nutrient absorption (Neumam et al., 2017NEUMAN, E.R.; RESENDE, J.T.; CAMARGO, L.K.P.; CHAGAS, R.R.; FILHO, R.B.L. Produção de mudas de batata-doce em ambiente protegido com aplicação de extrato de Ascophyllum nodosum. Horticultura Brasileira, v.35, n.2, p.490-498, 2017. http://dx.doi.org/10.1590/S0102-053620170404
http://dx.doi.org/10.1590/S0102-05362017...
).

Another bioregulator used is Stimulate®, whose function is to establish hormonal balance, differentiation, cell elongation, and promote germination of seeds and better growth of the root system, promoting a greater capacity to absorb nutrients and water by the roots of higher plants (Rós et al., 2014RÓS, A.B.; NARITA, N.; ARAÚJO, H.S. Efeito de bioestimulante no crescimento inicial e na produtividade de plantas de batata-doce. Revista Ceres , v.62, n.5, p.469-474, 2015. https://doi.org/10.1590/0034-737X201562050007
https://doi.org/10.1590/0034-737X2015620...
; Smiderle and Souza, 2021SMIDERLE, O.J.; SOUZA, A.G. Do scarification and seed soaking periods promote maximum vigor in seedlings of Hymenaea courbaril?. Journal of Seed Science , v.43, p.e202143030, 2021. https://doi.org/10.1590/2317-1545v43254481
https://doi.org/10.1590/2317-1545v432544...
).

Trichoderma spp. can generate benefits in the processes of germination as well as in the initial growth of seedlings of forest species through the treatment of seeds and application in soil (Costa et al., 2018COSTA, N.C.R.; CARVALHO, F.J.; JULIATTI, F.C.; CASTRO, M.V.; CUNHA, W.V. Diversity of fungi in seeds of Hymenaea stignocarpa and Hymenaea courbaril before and after fungicide treatments. Bioscience Journal, v.34, n.4, p.868-874, 2018. https://doi.org/10.14393/BJ-v34n1a2018-41866
https://doi.org/10.14393/BJ-v34n1a2018-4...
; Junges et al., 2016JUNGES, E.; MUNIZ, M.F.; MEZZOMO, R.; BASTOS, B.; MACHADO, R.T. Trichoderma spp. na produção de mudas de espécies florestais.Floresta e Ambiente , v.23, n.2, p.237-244, 2016. https://dx.doi.org/10.1590/2179-8087.107614
https://dx.doi.org/10.1590/2179-8087.107...
). In addition, Junges et al. (2016)JUNGES, E.; MUNIZ, M.F.; MEZZOMO, R.; BASTOS, B.; MACHADO, R.T. Trichoderma spp. na produção de mudas de espécies florestais.Floresta e Ambiente , v.23, n.2, p.237-244, 2016. https://dx.doi.org/10.1590/2179-8087.107614
https://dx.doi.org/10.1590/2179-8087.107...
reported positive results in the gain of the percentage of emergence of Peltophorum dubium (Fabaceae) seedlings promoted by Trichoderma spp., pointing to its potential to be used in the treatment of orthodox seeds.

In view of the above, the present study establishes the following research problem: Can different concentrations of Acadian®, Stimulate®, and Trichoderma spp. be effective in promoting the emergence and vigor of H. courbaril seedlings with or without mechanical scarification?

MATERIAL AND METHODS

The study was conducted in the Seed Analysis Laboratory (LAS) and the seedling nursery of the forestry sector of Embrapa Roraima from May to October 2020. The species used in the present study was H. courbaril. Its fruits were hand-collected from trees present in an area of Submontane Dense Ombrophilous Forest with emerging canopy, located at the geographic coordinates 1°38’29” North latitude and 60°58’11” West longitude, in the municipality of Boa Vista (RR), Brazil, in April 2020.

The climate of the municipality of Boa Vista - RR is Am (tropical monsoon climate), with average temperatures of 27.2 °C in the hottest month and 23.3 °C in the coldest month, with an annual average of 25.4 °C. The average annual rainfall is 1808 mm, with mean values of 365 mm and 26 mm for the months of highest (June) and lowest (February) precipitation, respectively.

The collected ripe fruits were selected by removing from the lot those with mechanical injury and deteriorated, to obtain a uniform sample. Subsequently, they were properly cleaned, pulped and the seeds were washed in running water. For the biometric characterization of the seeds, they were measured in the middle region, using a digital caliper with an accuracy of 0.01. For the studies of pre-germination treatments, H. courbaril seeds were subjected to the mechanical scarification treatment, with sandpaper no 100, by manually frictioning them until the rupture of the seed coat and appearance of the cotyledonary tissues on the opposite side of the hilum/embryonic axis. The concentrations of Acadian®, Stimulate®, and Trichoderma spp. used were: 0.0, 5.0, 10.0, 15.0 mL.L-1 for each 0.5 kg of seeds, applied as pre-germination treatment by soaking the seeds for 30 minutes. Seeds that did not receive bioregulators served as a control. Seed treatment was carried out according to the methodology recommended by Nunes (2005NUNES, J.C. Tratamento de semente - qualidade e fatores que podem afetar a sua performance em laboratório. Syngenta Proteção de Cultivos Ltda. 2005. 16p.), in which the different concentrations of the bioregulators were applied directly to the bottom of a transparent plastic bag for each treatment, and then the seeds were placed inside the plastic bag and shaken manually for 30 minutes so that the hormone reached all seeds.

The experimental design used was completely randomized, in a 4 x 3 x 2 factorial scheme, corresponding to four concentrations (0, 5.0, 10.0 and 15.0 mL.L-1), three hormones (Stimulate®, Acadian® and Trichoderma spp.) with and without mechanical scarification of seeds, with four replicates, each of which consisting of 25 seeds.

In order to complement and elucidate the results of the present study, the seeds were sown in sand of medium particle size at 1.0 cm depth in 30 cm x 40 cm x 10 cm plastic trays in a greenhouse with an average temperature of 27 ± 5 °C and relative humidity of 60% to 70% in the experimental period. The variables evaluated were: emergence speed (ES, index) (Maguire, 1962MAGUIRE, J.D. Speed of germination-aid in selection and evaluation for seedling emergence and vigor. Crop Science, v.2, n.1, p.176-177, 1962. https://doi.org/10.2135/cropsci1962.0011183X000200020033x
https://doi.org/10.2135/cropsci1962.0011...
) and seedling emergence (E, %), obtained by daily counts until the stabilization of the counts of all treatments. During the initial vigor stages of the seedlings, substrate moisture content was maintained by manual irrigation, in a total of four daily irrigations. Seedling emergence started nine days after sowing (DAS).

Results of vigor were expressed as a percentage of normal seedlings, counted during the evaluations. A normally emerged seedling was the one which, after breaking the surface of the substrate, showed well-differentiated and developed plumule, cotyledons, and hypocotyl.

To verify the assumptions of the analysis of variance (ANOVA), the data were first assessed for: a) normality with Shapiro-Wilk test (p<0.01) and b) homoscedasticity by Bartlett test (p<0.01). When there was the normality of residuals and homogeneity of variances, the data were subjected to ANOVA, followed by Tukey test (p<0.01) to compare the means. Quantitative variables were subjected to regression analysis to verify the response of seed vigor as a function of the concentrations. Data analysis was performed in the statistical program Sisvar (Ferreira, 2014FERREIRA, D.F. Sisvar: a Guide for its Bootstrap procedures in multiple comparisons. Ciência e Agrotecnologia, v.38, n.2, p.109-112, 2014.https://doi.org/10.1590/S1413-70542014000200001
https://doi.org/10.1590/S1413-7054201400...
).

RESULTS AND DISCUSSION

For the biometric characterization of the seeds, their mean values were 26.38 (length (mm)), 20.57 (width (mm)), and 12.37 (thickness (mm)) and individual fresh mass of H. courbaril seeds (mean value of 5.70 g). Similar values were described by Andrade et al. (2010ANDRADE, L.A.; BRUNO, R.D.L.A.; LUCENA, R.; OLIVEIRA, L.S.B.; SILVA, H.T.F. Aspectos biométricos de frutos e sementes, grau de umidade e superação de dormência de jatobá. Acta Scientiarum Agronomy, v.32, n.2, p.293-299, 2010. http://www.scielo.br/pdf/asagr/v32n2/a16v32n2.pdf
http://www.scielo.br/pdf/asagr/v32n2/a16...
) and Duarte et al. (2016DUARTE, M.M.; PAULA, S.R.P.; FERREIRA, F.R.L.; NOGUEIRA, A.C. Morphological characterization of fruit, seed and seedling and germination of Hymenaea courbaril L. (Fabaceae) (‘Jatobá’). Journal of Seed Science, v.38, n.3, p.204-211, 2016. https://doi.org/10.1590/2317-1545v38n3159734
https://doi.org/10.1590/2317-1545v38n315...
) for Hymenaea courbaril, by Botelho et al. (2000BOTELHO, S.A.; FERREIRA, R.A.; MALAVASI, M.M.; DAVIDE, A.C. Aspectos morfológicos de frutos, sementes, plântulas e mudas de jatobá do-cerrado (Hymenaea stigonocarpa Mart. ex Hayne)-Fabaceae. Revista Brasileira de Sementes, v.22, n.1, p.144-152, 2000. ) for Hymenaea stigonocarpa Mart.ex Hayne and Cruz et al. (2001CRUZ, E.D.; MARTINS, F.O.; CARVALHO, J.E.U. Biometria de frutos e sementes de jatobá-curuba (Hymenaea intermedia Ducke, Leguminosae - Caesalpinioideae). Revista Brasileira de Botânica, v.24, n.2, p.161-165, 2001. https://doi.org/10.1590/S0100-84042001000200005
https://doi.org/10.1590/S0100-8404200100...
) for Hymenaea intermedia.

According to the analysis of variance, there was no significant interaction between the factors: bioregulator (B), Scarification (S), and Concentrations (C), for the variables seedling emergence (%E), emergence speed (ES, index), and normal seedling (NS) (Table 1); therefore, the effect of each factor was evaluated separately. In addition, it showed a significant effect of factors (B), (S) and (C), isolated, for the variables seedling emergence (%E), emergence speed (ES, index) and normal seedling (NS) of H. courbaril.

Table 1
Summary of the analysis of variance for seedling emergence (%E), emergence speed (ES, index) and normal seedling (NS) of H. courbaril as a function of the bioregulator (B), concentrations (0, 5, 10 and 15 mL.L-1) and physical scarification (with and without), as well as their interactions.

H. courbaril seeds showed higher results for seedling emergence percentage, in the presence of mechanical scarification at concentration of 0.0 mL.L-1, i.e., control (Table 1), compared to non-scarified seeds (control), suggesting that scarification with sandpaper no 100 on the opposite side of the hilum/embryonic axis is adequate for overcoming seed dormancy and inducing emergence.

These results corroborate those reported by Costa et al. (2017COSTA, C.H.M.; DIARIS, K.B.; GUIMARÃES, T.M. Metodos de escarificação para superação de dormência de sementes de jatobá. Revista Científica Eletrônica de Engenharia Florestal, v.30, n.1, p.44-52, 2017.) for H. courbaril in a study on the effect of scarification with sandpaper no 100 on the opposite side of the hilum/embryonic axis, as these authors also verified that, under such conditions, it was possible to obtain superiority and uniformity in the emergence of seedlings of this species.

In general, in the present study, the emergence of seedlings, with seeds of H. courbaril without mechanical scarification, obtained mean values of 48.5%, results considered unsatisfactory according to Brasil (2009). Ribeiro et al. (2021RIBEIRO, O.D.; CRUZ, E.D.; SILVA, M.F.; CHAVES, B.A.; RIBEIRO, O.M.D.; GURGEL, E.S.C. Hymenaea parvifolia Huber: dormancy breaking, morphology of fruit, seed and seedling. Revista Ceres, v.68, n.2, p.105-114, 2021. https://doi.org/10.1590/0034-737X202168020003
https://doi.org/10.1590/0034-737X2021680...
), evaluating different methods of scarification in seeds of this same species, recorded zero seedling emergence in seeds without scarification (control). These results make it possible to indicate with reliability the use of the pre-germination treatment of mechanical scarification to overcome the physical dormancy of H. courbaril seeds.

Figure 1 (A) showed that the percentage of emergence of H. courbaril seedlings in relation to the concentrations attributed to the bioregulator Acadian® showed that the concentrations of 5.0 and 10.0 mL.L-1 with mechanical scarification resulted in a 12% increase in seedling emergence when compared with seeds that did not receive the bioregulator (Figure 1 A). In turn, seeds without mechanical scarification and Acadian® concentrations showed a decrease of 34% in emergence compared to those with scarification and in the absence of Acadian® concentrations for the same variable (Figure 1 A). In addition, there was a 33% superiority in the emergence of H. courbaril seedlings without mechanical scarification of seeds at the concentration 10 mL.L-1 of Acadian®, which led to similar performance to the results obtained with seeds scarified at zero concentration, that is, in the absence of the bioregulator Acadian® (Figure 1 B).

Figure 1
Mean values of seedlings emergence obtained using three bioregulators Acadian® (A), Stimulate® (B) Trichoderma spp. (C) at four concentrations 0, 5.0, 10.0 and 15.0 mL.L-1 respectively, and as well as a table (D) with Tukey test results in H. courbaril seeds without and with mechanical scarification.

In addition, for the emergence of H. courbaril seedlings the estimated dose of maximum technical efficiency (DMTE) of Acadian® for seeds with scarification was 8.50 mL.L-1, corresponding to 86.09% seedling emergence (Figure 1), whereas for seeds without scarification the DMTE was 10.10 mL.L-1 of Acadian®, which resulted in 75.21% seedling emergence (Figure 1).

In turn, Stimulate® led to seedling emergence of approximately 90% at concentration of 10.0 mL.L-1, which was also found with 15.0 mL.L-1 of the bioregulator (Figure 1 B). However, for the emergence of seedlings with the bioregulator Stimulate®, the estimated DMTE was 9.17 mL.L-1 for seeds without mechanical scarification, which led to emergence percentage of 75.21% (Figure 1B).

It is suggested that this interaction between hormone and Stimulate® concentrations can enhance the maintenance of physiological processes that culminate with cytokinin activity and influences cell elasticity and plasticity (growth), promoting different responses in the vigor of H. courbaril seedlings (Kolachevskaya et al., 2017KOLACHEVSKAYA, O.O.; SERGEEVA, L.; FLOKOVA, K.; GETMAN, I.A.; LOMIN, S.N.; ALEKSEEVA, V.V. Auxin synthesis gene tms1 driven by tuberspecific promoter alters hormonal status of transgenic potato plants and their responses to exogenous phytohormones. Plant Cell Reports, v.36, n.2, p.419-435, 2017. https://doi.org/10.1007/s00299-016-2091-y
https://doi.org/10.1007/s00299-016-2091-...
).

In addition, the emergence of seedlings with mechanical scarification of seeds without Stimulate®, compared with non-scarified seeds, was 30% lower, a result that confirms the inhibitory action of the seed coat on this variable (Figure 1 B).

The results of the present study reinforce the importance of knowing the emergence potential of H. courbaril seedlings native to the northern region of Brazil, for commercial use by producers and nursery managers in the state of Roraima.

Although the concentration of 5.0 mL.L-1 of Trichoderma spp. applied to H. courbaril seeds with mechanical scarification resulted in the maximum percentage of emergence (75%), with DMTE of 7.17 mL.L-1 promoting 76.45%, for the Stimulate® hormone the maximum percentage of emergence (90%) was obtained at a concentration of 10.0 mL.L-1 in seeds with mechanical scarification. Furthermore, the concentrations of 5.0 and 10.0 mL.L-1 of the bioregulator obtained from algae extracts (Acadian®) applied in H. courbaril seeds with mechanical scarification resulted in 85% seedling emergence (Figure 1 A). These results make it possible to indicate with greater reliability that it is feasible to perform pre-germination treatment using mechanical scarification in H. courbaril seeds and the aforementioned concentrations of plant bioregulators (Acadian®, Stimulate® and Trichoderma spp.) to overcome seed dormancy, aiming at large-scale production of seedlings.

When comparing the emergence speed of H. courbaril seedlings with and without mechanical scarification of seeds between the concentrations of the three plant bioregulators tested, it was found that H. courbaril seeds with mechanical scarification led to higher ES (index) at all concentrations of bioregulators tested compared to the control, except for the concentration of 15.0 mL.L-1 of Trichoderma spp., as shown in Figures 2 A, B, C.

Figure 2
Mean values for seedling emergence speed obtained with treatment using three bioregulators Acadian® (A), Stimulate® (B) Trichoderma spp. (C) at four concentrations 0, 5.0, 10.0 and 15.0 mL.L-1 respectively, and as well as a table (D) with Tukey test results in H. courbaril seeds without and with mechanical scarification.

In turn, the DMTE for emergence speed of H. courbaril seedlings with mechanical scarification (Figure 2 A) was obtained with the concentration of 14.29 mL.L-1 of the hormone Stimulate® with 6.86 ES (index). There was also superiority in emergence speed with seeds subjected to mechanical scarification and treated with the hormone Acadian® at a DMTE of 9.30 mL.L-1 with 7.16 ES (index), which led to values close to the results obtained with Stimulate® at a concentration of 14.29 mL.L-1 in scarified seeds (Figures 2 A, B).

The results of the present study showed that appropriate concentrations of Stimulate®, Acadian®, and Trichoderma spp. positively influenced the emergence of H. courbaril seedlings, except for the concentration of 15.0 mL.L-1 of Trichoderma spp., which may have been excessive, even causing inhibition. According to Souza et al. (2017SOUZA, A.G.; SMIDERLE, O.J.; SPINELLI, V.M.; SOUZA, R.O.; BIANCHI, V.J. Optimization of germination and initial quality of seedlings of Prunus persica tree rootstocks. Journal of Seed Science , v.39, n.2, p.166-173, 2017. http://dx.doi.org/10.1590/2317-1545v39n2171687
http://dx.doi.org/10.1590/2317-1545v39n2...
), a faster germination (higher ES [index]) stay for lower time subject to adverse conditions found in the soil, such as temperature variation, water stress, and attack of pests and pathogens. In this case, the use of plant bioregulators was favorable to the emergence speed of H. courbaril seedlings, being indicated under the conditions of the present study.

The results obtained in this study reinforced those reported by Carvalho et al. (2018CARVALHO, J.H.N.; LIMA, A.P.L.; LIMA, S.F. Adição de moinha de carvão e de Stimulate® na formação de mudas de Acacia mangium. Revista de Agricultura Neotropical, v.5, n.1, p.66-74, 2018.), who found that the ideal concentration of Stimulate® optimized the percentage of emergence in Acacia mangium (Fabaceae), which has orthodox seeds. Oliveira et al. (2020OLIVEIRA, E.; OLIVEIRA, J. C.; FILIPINI, T.O.; SOUZA, G.C.; VILÃO, A.C.; SOUZA, F.M.L. Superação de dormência em sementes de (Hymenaea courbaril) com regulador e estimulante vegetal. Revista Científica Eletrônica de Ciências Aplicadas, v.15, n.1, p.1-7, 2020.) verified that H. courbaril seeds without mechanical scarification and treated with Stimulate® at concentrations of 20 and 30 mL.L-1 had values lower than 24% in the percentage of emergence, and these concentrations of Stimulate® contributed decisively to the inhibitory or reductional effect on this variable. According to Singh et al. (2019SINGH, V.; SERGEEVA, L.; LIGTERINK, W.; ALONI, R.; ZEMACH, H.; DORON, F.A; YANG, J.; ZHANG, P.; SHABTAI, S.; FIRON, N. Gibberellin promotes sweetpotato root vascular lignification and reduces storage-root formation. Frontiers Plant Science, v.10, n.3, p.1320, 2019. doi: 10.3389/fpls.2019.01320
https://doi.org/10.3389/fpls.2019.01320...
), inadequate concentrations of plant bioregulators can lead to deterioration due to phytosanitary, physiological, biochemical, and cytological alterations in the seeds, culminating in low emergence or even embryo death.

In mechanical scarification with the presence of the bioregulator Acadian® at a concentration of 5.0 mL.L-1, on average the emergence speed (index) was 27 percentage points higher when compared with Trichoderma spp. at a concentration of 5.0 mL.L-1, while the highest value of emergence speed (6.84) for Stimulate® was found at a concentration of 10.0 mL.L-1 with mechanical scarification (Figures 2 A, B, C). Thus, it demonstrates that the mechanical scarification of H. courbaril seeds concomitantly with hormone and the above-mentioned concentrations allow H. courbaril seeds to express maximum vigor and establish vigorous seedlings (Figures 3 A, B, C).

Figure 3
Mean values of the percentage of normal seedlings obtained with the treatment using three bioregulators Acadian® (A), Stimulate® (B) Trichoderma spp (C) at four concentrations 0, 5.0, 10.0 and 15.0 mL.L-1 respectively, and as well as a table (D) with Tukey test results in H. courbaril seeds without and with mechanical scarification.

In addition, the highest percentage of normal seedlings (85%) was found with DMTE of 8.83 mL.L-1 of Acadian® in scarified seeds, while for non-scarified seeds the DMTE of Acadian® was 10.0 mL.L-1 and it led to a value of 70%, that is, there was a 17.65 percentage points increase to obtain normal H. courbaril seedlings.

On the other hand, for H. courbaril seeds without mechanical scarification and without the addition of Acadian®, the percentage of normal seedlings was equal to 50% (Figure 3 A), suggesting once again the need for mechanical scarification for the continuity of the germination process by the embryo. The results described in the present study as well as in the national and international literature show that the physical dormancy of H. courbaril seeds is always present (Pierezan et al., 2012PIEREZAN, L., SCALON, S.P.Q., PEREIRA, Z.V. Emergência de plântulas e crescimento de mudas de jatobá com uso de bioestimulante e sombreamento. Cerne, v.18, n.2, p.127- 133, 2012. https://doi.org/10.1590/S0104-77602012000100015.
https://doi.org/10.1590/S0104-7760201200...
; Soares et al., 2017SOARES, L.V.; OLIVEIRA, S.C.C.; BRAGA, L.R.; SAMPAIO, A.B.; SCHMIDT, I.B. Can phytohormones stimulate initial growth of brazilian savanna trees? Heringeriana, v.11, n.2, p.1-12, 2017. https://doi.org/10.17648/heringeriana.v11i2.917736
https://doi.org/10.17648/heringeriana.v1...
; Costa et al., 2019COSTA, E.; LOPES, K.G.; BINOTTI, F.F.S.; CARDOSO, E.D.C.; DALASTRA, C. Technologies for jatobá seedling formation.Floresta e Ambiente, v.26, n.1, p.e20150084, 2019.https://doi.org/10.1590/2179-8087.008415
https://doi.org/10.1590/2179-8087.008415...
; Dourado et al., 2020DOURADO, D.; LIMA, S.F.; LIMA, A.P.; SORATTO, D.N.; BERNARDO, V.F.; BARBOSA, H.M. Efeito de bioestimulante em sementes de cedro-rosa. Brazilian Journal of Development, v.6, n.5, p.30306-30319, 2020. https://www.brazilianjournals.com/index.php/BRJD/article/view/10503
https://www.brazilianjournals.com/index....
) and needs to be overcome with mechanical scarification and as well as adequate concentrations of hormones based on cytokinin, indole butyric acid and gibberellic acid applied in the seeds.

Additionally, the presence of the lignified coat in the seeds may limit the vigor of H. courbaril seedlings, resulting in an uneven stand of seedlings, confirmed by the works of Dourado et al. (2020DOURADO, D.; LIMA, S.F.; LIMA, A.P.; SORATTO, D.N.; BERNARDO, V.F.; BARBOSA, H.M. Efeito de bioestimulante em sementes de cedro-rosa. Brazilian Journal of Development, v.6, n.5, p.30306-30319, 2020. https://www.brazilianjournals.com/index.php/BRJD/article/view/10503
https://www.brazilianjournals.com/index....
) and Costa et al. (2019COSTA, E.; LOPES, K.G.; BINOTTI, F.F.S.; CARDOSO, E.D.C.; DALASTRA, C. Technologies for jatobá seedling formation.Floresta e Ambiente, v.26, n.1, p.e20150084, 2019.https://doi.org/10.1590/2179-8087.008415
https://doi.org/10.1590/2179-8087.008415...
) and also demonstrated with the data of the present study.

An increase in the percentage of normal seedlings according to the increase in the DMTE of 11.83 mL.L-1 with 84.91% of Stimulate® was found in scarified seeds (Figure 3 B), indicating that the input of Stimulate® solution was sufficient to promote effective changes in hormonal balance.

On the other hand, it was verified that seeds without mechanical scarification at the concentration of 10.0 mL.L-1 of Stimulate® led to 70% of normal seedlings, while the concentration of 5.0 mL.L-1 of Stimulate® resulted in 65% of normal seedlings and the concentration of 15.0 mL.L-1 of Stimulate® promoted 60% of normal seedlings.

On the other hand, seeds without mechanical scarification and without the addition of Stimulate® (Figure 3 B) led to the lowest percentage of normal seedlings (45%) compared to the Stimulate® concentrations mentioned above.

These results show that the rigid coat, involving the seeds of the species under study, does not totally restrict the entry of water and hormones to overcome dormancy, but probably the differentiated concentrations of Stimulate® were sufficient to promote significant changes in the hormonal balance that controls the physiological dormancy process, making the metabolism of seeds more active and the embryo fit for resuming the development and consequently promoting more gain of normal H. courbaril seedlings.

According to Zerlin et al. (2016ZERLIN, J.K.; CENTENO, C.; GASPAR, M. Exogenous nitric oxide-induced germination associated with pinitol content inHymenaea courbarillow vigour seeds.Brazilian Journal of Botany, v.39, v.5, p.485-494, 2016. https://doi.org/10.1007/s40415-016-0265-z
https://doi.org/10.1007/s40415-016-0265-...
), gibberellin at concentrations suitable for each species, when applied exogenously, can double the synthesis of proteins in seeds, which favors germination. However, the promotion of gains in seedling vigor for all studied variables was verified in the present study for H. courbaril seeds.

Souza et al. (2016SOUZA, A.G.; SMIDERLE, O.J.; SPINELLI, V.M.; SOUZA, R.O.; BIANCHI, V.J. Correlation of biometrical characteristics of fruit and seed with twinning and vigor of Prunus persica rootstocks. Journal of Seed Science , v.38, n.4, p.322-328, 2016. http://dx.doi.org/10.1590/2317-1545v38n4164650
http://dx.doi.org/10.1590/2317-1545v38n4...
) reported that, during the period of stratification of Prunus persica seeds at 7 °C, there was an increase in the production of normal seedlings due to the adequate concentration of growth-promoting substances, such as gibberellins and indoleacetic acid, as well as the reduction of abscisic acid, a germination-inhibiting substance. According to Menegatti et al. (2019MENEGATTI, R.D.; SOUZA, A.G.; BIANCHI, V.J. Estimating genetic divergence between peach rootstock cultivars using multivariate techniques based on characteristics associated with seeds. Genetics and Molecular Research, v.1, n.3, p.01-10, 2019. http://dx.doi.org/10.4238/gmr18345
http://dx.doi.org/10.4238/gmr18345...
), a fully developed embryo requires specific and adequate conditions to promote complete development and ensure the establishment of a new seedling.

Figure 3 C shows that seeds with mechanical scarification and inoculated with Trichoderma spp. at the DMTE of 5.50 mL.L-1 led to 65.0% of normal seedlings, compared with the other concentrations. According to Normative Instruction 45 (IN 45), which regulates the production of seeds from commercial crops in Brazil, the minimum germination acceptable for commercialization is 80%.

Therefore, considering the recommended standard and according to the results obtained, it is possible to infer that seeds with and without scarification and concentrations of Trichoderma spp. did not reach the minimum value of emergence stipulated by the legislation, generally showing 50 to 65% of normal seedlings.

The results of the present study reinforce the importance of knowing the performance of Hymenaea courbaril seeds from Roraima state and combinations of bioregulator concentrations regarding seedling vigor, which can provide information for the selection of plant material suitable for commercial use by producers and nursery managers in the microregion of Boa Vista-Roraima.

A higher percentage of vigorous seedlings with adequate commercial characteristics is one of the primary factors for the success of the species, resulting in the economic return of the capital invested by producers and nursery managers.

It is known that a longer time of permanence of the seedlings in the nursery and the use of a greater number of seeds to compensate for the loss of viability and vigor generate a high cost of production. In addition, physical scarification on the opposite side of the hilum/embryonic axis of H. courbaril seeds requires labor and extra cost for the nursery manager.

However, this management is justified as it compensates for the loss in viability and vigor of seedlings. In addition, the physical scarification and treatments in seeds with the dose of maximum technical efficiency using plant bioregulators recommended in the present study for H. courbaril promote gains in the percentage of emergence, homogeneity and vigor of seedlings and, ultimately, reduction of costs and in the period to obtain commercial seedlings.

CONCLUSIONS

Acadian® at a concentration of 5.0 mL.L-1 in scarified seeds of H. courbaril leads to a greater increase in the percentage of seedling emergence and the percentage of normal seedlings.

The dose of maximum technical efficiency (DMTE) of the bioregulator Acadian® is 8.50 mL.L-1 and, in scarified seeds of H. courbaril, it increases the percentage of seedling emergence.

The dose of maximum technical efficiency (DMTE) of Stimulate® is 9.17 mL.L-1 to obtain a high percentage of emergence of H. courbaril seedlings with seeds subjected to mechanical scarification.

Trichoderma spp. from 9.0 mL.L-1 does not favor gains in the percentage of normal seedlings of H. courbaril grown from seeds with and without mechanical scarification.

Mechanical scarification of H. courbaril seeds with addition of treatments using plant bioregulators enables the efficient overcoming of physical and physiological dormancy, making it possible to optimize the rate of seedling emergence for large-scale production of uniform seedlings.

ACKNOWLEDGMENTS

The authors thank the Conselho Nacional de Desenvolvimento Científico eTecnológico (CNPq), for the productivity and research scholarships granted to the first author.

REFERENCES

  • ALMEIDA, M.B.; SOUZA, W.C.O.; GOMES, E.C.S.; RAMALHO, F.C.V. Descrição morfológica do fruto e semente do jatobá (Hymenaea courbaril L). Revista Semiárido de Visu, v.1, n.2, p.107-115, 2011. https://periodicos.ifsertao-pe.edu.br/ojs2/index.php/semiaridodevisu/article/view/39
    » https://periodicos.ifsertao-pe.edu.br/ojs2/index.php/semiaridodevisu/article/view/39
  • ANDRADE, L.A.; BRUNO, R.D.L.A.; LUCENA, R.; OLIVEIRA, L.S.B.; SILVA, H.T.F. Aspectos biométricos de frutos e sementes, grau de umidade e superação de dormência de jatobá. Acta Scientiarum Agronomy, v.32, n.2, p.293-299, 2010. http://www.scielo.br/pdf/asagr/v32n2/a16v32n2.pdf
    » http://www.scielo.br/pdf/asagr/v32n2/a16v32n2.pdf
  • BOTELHO, S.A.; FERREIRA, R.A.; MALAVASI, M.M.; DAVIDE, A.C. Aspectos morfológicos de frutos, sementes, plântulas e mudas de jatobá do-cerrado (Hymenaea stigonocarpa Mart. ex Hayne)-Fabaceae. Revista Brasileira de Sementes, v.22, n.1, p.144-152, 2000.
  • CARVALHO, J.H.N.; LIMA, A.P.L.; LIMA, S.F. Adição de moinha de carvão e de Stimulate® na formação de mudas de Acacia mangium Revista de Agricultura Neotropical, v.5, n.1, p.66-74, 2018.
  • COSTA, C.H.M.; DIARIS, K.B.; GUIMARÃES, T.M. Metodos de escarificação para superação de dormência de sementes de jatobá. Revista Científica Eletrônica de Engenharia Florestal, v.30, n.1, p.44-52, 2017.
  • COSTA, E.; LOPES, K.G.; BINOTTI, F.F.S.; CARDOSO, E.D.C.; DALASTRA, C. Technologies for jatobá seedling formation.Floresta e Ambiente, v.26, n.1, p.e20150084, 2019.https://doi.org/10.1590/2179-8087.008415
    » https://doi.org/10.1590/2179-8087.008415
  • COSTA, N.C.R.; CARVALHO, F.J.; JULIATTI, F.C.; CASTRO, M.V.; CUNHA, W.V. Diversity of fungi in seeds of Hymenaea stignocarpa and Hymenaea courbaril before and after fungicide treatments. Bioscience Journal, v.34, n.4, p.868-874, 2018. https://doi.org/10.14393/BJ-v34n1a2018-41866
    » https://doi.org/10.14393/BJ-v34n1a2018-41866
  • CRUZ, E.D.; MARTINS, F.O.; CARVALHO, J.E.U. Biometria de frutos e sementes de jatobá-curuba (Hymenaea intermedia Ducke, Leguminosae - Caesalpinioideae). Revista Brasileira de Botânica, v.24, n.2, p.161-165, 2001. https://doi.org/10.1590/S0100-84042001000200005
    » https://doi.org/10.1590/S0100-84042001000200005
  • DOURADO, D.; LIMA, S.F.; LIMA, A.P.; SORATTO, D.N.; BERNARDO, V.F.; BARBOSA, H.M. Efeito de bioestimulante em sementes de cedro-rosa. Brazilian Journal of Development, v.6, n.5, p.30306-30319, 2020. https://www.brazilianjournals.com/index.php/BRJD/article/view/10503
    » https://www.brazilianjournals.com/index.php/BRJD/article/view/10503
  • DUARTE, M.M.; PAULA, S.R.P.; FERREIRA, F.R.L.; NOGUEIRA, A.C. Morphological characterization of fruit, seed and seedling and germination of Hymenaea courbaril L. (Fabaceae) (‘Jatobá’). Journal of Seed Science, v.38, n.3, p.204-211, 2016. https://doi.org/10.1590/2317-1545v38n3159734
    » https://doi.org/10.1590/2317-1545v38n3159734
  • FERREIRA, D.F. Sisvar: a Guide for its Bootstrap procedures in multiple comparisons. Ciência e Agrotecnologia, v.38, n.2, p.109-112, 2014.https://doi.org/10.1590/S1413-70542014000200001
    » https://doi.org/10.1590/S1413-70542014000200001
  • JUNGES, E.; MUNIZ, M.F.; MEZZOMO, R.; BASTOS, B.; MACHADO, R.T. Trichoderma spp. na produção de mudas de espécies florestais.Floresta e Ambiente , v.23, n.2, p.237-244, 2016. https://dx.doi.org/10.1590/2179-8087.107614
    » https://dx.doi.org/10.1590/2179-8087.107614
  • KOLACHEVSKAYA, O.O.; SERGEEVA, L.; FLOKOVA, K.; GETMAN, I.A.; LOMIN, S.N.; ALEKSEEVA, V.V. Auxin synthesis gene tms1 driven by tuberspecific promoter alters hormonal status of transgenic potato plants and their responses to exogenous phytohormones. Plant Cell Reports, v.36, n.2, p.419-435, 2017. https://doi.org/10.1007/s00299-016-2091-y
    » https://doi.org/10.1007/s00299-016-2091-y
  • MAGUIRE, J.D. Speed of germination-aid in selection and evaluation for seedling emergence and vigor. Crop Science, v.2, n.1, p.176-177, 1962. https://doi.org/10.2135/cropsci1962.0011183X000200020033x
    » https://doi.org/10.2135/cropsci1962.0011183X000200020033x
  • MENEGATTI, R.D.; SOUZA, A.G.; BIANCHI, V.J. Estimating genetic divergence between peach rootstock cultivars using multivariate techniques based on characteristics associated with seeds. Genetics and Molecular Research, v.1, n.3, p.01-10, 2019. http://dx.doi.org/10.4238/gmr18345
    » http://dx.doi.org/10.4238/gmr18345
  • MENEZES-FILHO, A.C.P.; SILVA, M.A.; PEREIRA, A.V.; OLIVEIRA FILHO, J. G.; CASTRO, C.F.S. Parâmetros físico-químicos, tecnológicos, atividade antioxidante, conteúdo de fenólicos totais e carotenóides das farinhas dos frutos do jatobá-do-cerrado (Hymenaea stigonocarpa Mart. ex Hayne). Multi-Science Journal, v.2, n.1, p.93-100, 2019. doi: 10.33837/msj.v2i1.900
    » https://doi.org/10.33837/msj.v2i1.900
  • NEUMAN, E.R.; RESENDE, J.T.; CAMARGO, L.K.P.; CHAGAS, R.R.; FILHO, R.B.L. Produção de mudas de batata-doce em ambiente protegido com aplicação de extrato de Ascophyllum nodosum Horticultura Brasileira, v.35, n.2, p.490-498, 2017. http://dx.doi.org/10.1590/S0102-053620170404
    » http://dx.doi.org/10.1590/S0102-053620170404
  • NUNES, J.C. Tratamento de semente - qualidade e fatores que podem afetar a sua performance em laboratório. Syngenta Proteção de Cultivos Ltda. 2005. 16p.
  • OLIVEIRA, E.; OLIVEIRA, J. C.; FILIPINI, T.O.; SOUZA, G.C.; VILÃO, A.C.; SOUZA, F.M.L. Superação de dormência em sementes de (Hymenaea courbaril) com regulador e estimulante vegetal. Revista Científica Eletrônica de Ciências Aplicadas, v.15, n.1, p.1-7, 2020.
  • PAIXÃO, M.V.S.; VIEIRA, K.M.; FERREIRA, E.A.; MÔNICO, A.F.; CARVALHO, A.J.C. Germination and dormity in jatobá seeds. International Journal of Advanced Engineering Research and Science, v.6, n.6, p.454-457, 2019. http://journal-repository.com/index.php/ijaers/article/view/578
    » http://journal-repository.com/index.php/ijaers/article/view/578
  • PIEREZAN, L., SCALON, S.P.Q., PEREIRA, Z.V. Emergência de plântulas e crescimento de mudas de jatobá com uso de bioestimulante e sombreamento. Cerne, v.18, n.2, p.127- 133, 2012. https://doi.org/10.1590/S0104-77602012000100015
    » https://doi.org/10.1590/S0104-77602012000100015
  • RAMOS, F.S.A.R.; SANTOS, T.C.; FERREIRA, T.H.B.; GOMES, M.C.S.; MUNHOZ, C.L. Aceitabilidade de biscoito tipo cookie enriquecidos com farinha de jatobá. Cadernos de Agroecologia, v.13, n.2, p.1-7, 2018. http://cadernos.aba-agroecologia.org.br/index.php/cadernos/article/view/2311/2157
    » http://cadernos.aba-agroecologia.org.br/index.php/cadernos/article/view/2311/2157
  • RIBEIRO, O.D.; CRUZ, E.D.; SILVA, M.F.; CHAVES, B.A.; RIBEIRO, O.M.D.; GURGEL, E.S.C. Hymenaea parvifolia Huber: dormancy breaking, morphology of fruit, seed and seedling. Revista Ceres, v.68, n.2, p.105-114, 2021. https://doi.org/10.1590/0034-737X202168020003
    » https://doi.org/10.1590/0034-737X202168020003
  • RISTAU, A.C.P.; CRUZ, M.S.F.V.; REGO, C.A.R.M.; BRAZ, H.; OLIVEIRA, S.S.; COSTA, P.B.; MALAVASI, M.M.; MALAVI, C.U.; TSUTSUMI, C.Y. Biometric characterisation and physiological quality of seeds of Hymenaea courbaril L. Journal of Experimental Agriculture International, v.26, n.3, p.1-9, 2018. https://doi.org/10.9734/JEAI/2018/43824
    » https://doi.org/10.9734/JEAI/2018/43824
  • RÓS, A.B.; NARITA, N.; ARAÚJO, H.S. Efeito de bioestimulante no crescimento inicial e na produtividade de plantas de batata-doce. Revista Ceres , v.62, n.5, p.469-474, 2015. https://doi.org/10.1590/0034-737X201562050007
    » https://doi.org/10.1590/0034-737X201562050007
  • SHANLEY, P.; MEDINA G. Frutíferas e plantas úteis na vida Amazônica- Jatobá Hymenaea courbaril Belém: CIFOR, Imazon, 2005. 300 p.
  • SILVA, C.P.; MANOLIO, S.F.R.A.; SAMPAIO, G.R.; BARROS, M.C.S.; NASCIMENTO, T.P.; CAMERON, L.C.; LARRAZ, M.S.F.; GOMES, J.A.A. Identification and action of phenolic compounds of Jatobá-do-cerrado (Hymenaea stignocarpa Mart.) on α-amylase and α-glucosidase activities and flour effect on glycemic response and nutritional quality of breads. Food Research International, v.116, n.18, p.1076-1083, 2019. https://pubmed.ncbi.nlm.nih.gov/30716891/
    » https://pubmed.ncbi.nlm.nih.gov/30716891/
  • SMIDERLE, O.J.; SOUZA, A.G. Do scarification and seed soaking periods promote maximum vigor in seedlings of Hymenaea courbaril?. Journal of Seed Science , v.43, p.e202143030, 2021. https://doi.org/10.1590/2317-1545v43254481
    » https://doi.org/10.1590/2317-1545v43254481
  • SINGH, V.; SERGEEVA, L.; LIGTERINK, W.; ALONI, R.; ZEMACH, H.; DORON, F.A; YANG, J.; ZHANG, P.; SHABTAI, S.; FIRON, N. Gibberellin promotes sweetpotato root vascular lignification and reduces storage-root formation. Frontiers Plant Science, v.10, n.3, p.1320, 2019. doi: 10.3389/fpls.2019.01320
    » https://doi.org/10.3389/fpls.2019.01320
  • SOARES, L.V.; OLIVEIRA, S.C.C.; BRAGA, L.R.; SAMPAIO, A.B.; SCHMIDT, I.B. Can phytohormones stimulate initial growth of brazilian savanna trees? Heringeriana, v.11, n.2, p.1-12, 2017. https://doi.org/10.17648/heringeriana.v11i2.917736
    » https://doi.org/10.17648/heringeriana.v11i2.917736
  • SOUZA, A.G.; SMIDERLE, O.J.; SPINELLI, V.M.; SOUZA, R.O.; BIANCHI, V.J. Correlation of biometrical characteristics of fruit and seed with twinning and vigor of Prunus persica rootstocks. Journal of Seed Science , v.38, n.4, p.322-328, 2016. http://dx.doi.org/10.1590/2317-1545v38n4164650
    » http://dx.doi.org/10.1590/2317-1545v38n4164650
  • SOUZA, A.G.; SMIDERLE, O.J.; SPINELLI, V.M.; SOUZA, R.O.; BIANCHI, V.J. Optimization of germination and initial quality of seedlings of Prunus persica tree rootstocks. Journal of Seed Science , v.39, n.2, p.166-173, 2017. http://dx.doi.org/10.1590/2317-1545v39n2171687
    » http://dx.doi.org/10.1590/2317-1545v39n2171687
  • ZERLIN, J.K.; CENTENO, C.; GASPAR, M. Exogenous nitric oxide-induced germination associated with pinitol content inHymenaea courbarillow vigour seeds.Brazilian Journal of Botany, v.39, v.5, p.485-494, 2016. https://doi.org/10.1007/s40415-016-0265-z
    » https://doi.org/10.1007/s40415-016-0265-z

Publication Dates

  • Publication in this collection
    15 Apr 2022
  • Date of issue
    2022

History

  • Received
    18 Mar 2021
  • Accepted
    15 Mar 2022
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