Impact of climate on seed morphology and plant growth of Caesalpinia bonduc L. in West Africa International Journal of Agronomy and Agricultural Research (IJAAR)

International Journal of Agronomy and Agricultural Research (IJAAR) ISSN: 2223-7054 (Print) 2225-3610 (Online) http://www.innspub.net Vol. 6, No. 3, p. 86-96, 2015 OPEN ACCESS RESEARCH PAPER Impact of climate on seed morphology and plant growth of Caesalpinia bonduc L. in West Africa Elie A. Padonou1*, Oscar D. Ahossou1, Farris O.Y. Okou1, Achille E. Assogbadjo1, Romain Glèlè Kakaï2, Anne Mette Lykke3, Brice Sinsin1 1 Laboratory of Applied Ecology, University of Abomey-Calavi, Cotonou, Benin Laboratory of Biomathematics and Forest Estimations, University of Abomey-Calavi, Cotonou, Benin 2 3 Department of Bioscience, Aarhus University, Silkeborg, Denmark Article published on March 21, 2015 Key words: Sudanian and Guinean climate zones, Hierarchical classification, Canonical discriminant analysis, Morphotypes. Abstract Caesalpinia bonduc L. is an important medicinal plant threatened by overexploitation. In the present study, the impact of climate on seed morphology, germination capacity, seedling and plant growth of C. bonduc were evaluated. A total of 2000 seeds were collected in Sudanian and Guinean climate zones of Africa and their length, width, thickness, weight and color were recorded. A hierarchical classification and canonical discriminant analysis were applied to the above traits of seeds from the different climatic zones. An analysis of variance with repeated measures was applied to seeds morphotypes identified by the hierarchical classification to test for the effect of these morphotypes on seed germination, seedling and plant growth. Hierarchical classification helped to identify four seed morphotypes. Canonical discriminant analysis performed on these morphotypes revealed highly significant differences. Morphotypes 1 and 3 comprised green seeds mainly from Sudanian zone while morphotypes 2 and 4 gathered grey seeds mainly from Guinean zone. Morphotype 3 had the longest seeds while the shortest seeds were from morphotype 1. The heaviest seeds were found in morphotype 4 whereas the lightest ones were from morphotype 1. Seeds of morphotype 4 were the thickest and widest, while the slimmest and most narrow ones were grouped in morphotype 1. Moprhotype 3, consisting of large green seeds mainly from Sudanian zone, was superior in terms of seedling and plant growth among all morphotypes and should be the best choice for planting purposes of the species. * Corresponding Author: Elie A. Padonou  padonouelie@yahoo.fr Padonou et al. Page 86 Introduction 2010; Padonou et al., 2013, 2014). Thus the present Caesalpinia bonduc L. is an important medicinal study aims to contribute to the conservation and plant widely distributed throughout the tropical and domestication of C. bonduc by (i) assessing the level subtropical regions of the world (Anonymous, 1956; of natural variation in seed morphology related to the Kapoor, 1990). Different parts of the plant are used to climatic zones in order to identify C. bonduc treat several diseases (Jain et al., 1992; Nandkarni, morphotypes based on seed characteristics (ii) 1976; Jethmalani et al., 1966), e.g. to ease childbirth, assessing the germination and plant growth ability of to treat burns and also for cultural practices like the identified morphotypes in order to determine the games, weddings and the Fâ ritual in Benin suitable (Assogbadjo et al., 2011). Different parts of the plant propagation of C. bonduc. morphotype and climate zone for have been found to have a variety of pharmacological activities (Simin et al., 2001; Kannur et al., 2006; Materials and methods Datté et al., 2004; Rastogi et al., 1996). The roots of Study area the plant are more intensively used than the leaves Seeds were collected in the Sudanian and Guinean and seed, which cause a stress to exploited climatic zones of Benin (Fig. 1). populations; moreover, there is low genetic diversity within the species which accentuates a need for conservation (Assogbadjo et al., 2012). The medicinal and sociocultural importance of the species had led to an overexploitation making it a rare and endangered species (Harden, 2002), and in Benin C. bonduc has been reported to be extinct in the wild, although it can be found in home gardens from the Guinean zone to the Sudanian zone (Adomou, 2005; Assogbadjo et al., 2012). Few researches have addressed the conservation and domestication of C. bonduc in Benin. The existing results were only related to the contribution of ethnic migrations on the propagation and persistence of the species and its morphological variability between climate zones (Assogbadjo et al 2012); the ethnic differences in use value and use patterns of the species (Assogbadjo et al 2011); and the germination technique of the seeds of the species (Hessou, 2009). Little is known about the morphological variation among seeds of C. bonduc in different climate zones although the local environment is known to shape morphological trait of seed (Salazar and Quesada, 1987; Assogbadjo et al., 2005; 2006). Moreover, the link between seed morphology, seed germination, and seedling and plant growth of C. bonduc were not yet established while many studies revealed a link between seed morphology, germination and plant growth (Assogbadjo et al., 2006 Fandohan et al., Figure 1. Climate zones of Benin The Sudanian zone is located between 9°45' and 12°25' N, while the Guinean zone is located between 6°25' and 7°30' N. The mean annual rainfall in the Sudanian zone is often less than 1000 mm and the relative humidity vary from 18 to 99% (highest in August). The temperature varies from 24 to 31°C. The Sudanian zone has hydromorphic, well-drained soils and lithosols. The vegetation in this zone is composed mainly of savannas with trees of smaller size. In the Guinean zone, the rainfall is bimodal with a mean annual rainfall of 1200 mm. The mean annual Padonou et al. Page 87 temperature varies between 25 and 29°C and the morphology. The same analysis was also performed to relative humidity ranges from 69 to 97%. The soils are test and describe the differences between the climate either deep ferralitic, or rich in clay, humus and zones according to seed morphology. minerals. The vegetation in this zone is mainly dense Germination, seedling and plant growth of C. semi-deciduous forest. bonduc seeds according to the morphotypes Identification and For germination tests on C. bonduc seeds and differences between the climate zones according to of measurement on seedling and plant growth, a nursery seed morphology experiment was carried out in January 2013 at the A total of 100 mature trees of C. bonduc, located at University of Abomey-Calavi, Benin (6°45’N; 2°35’ E least 100 m from each other, in order to avoid in the Guinean climatic zone). Four morphotypes of narrowing-down the genetic base due to relatedness seeds or inbreeding, were sampled in each climatic zone, as classification. For each morphotype, 90 seeds were recommended by Turnbull (1975). A total of 1000 sown; one in each pot (5.5 cm × 18 cm) made from a seeds were collected at random in each climatic zone. polythene bag and filled with forest soil. The seeds The collected seeds were kept under ambient were sown at an equal depth and the pots were temperature conditions for 10 days prior to sowing. watered equally twice daily (morning and evening) The length, width, and thickness of each seed were throughout the duration of the experiment. The measured using electronic calipers during these 10 experimental units were arranged in a randomized days. The weight of each seed was measured using complete block design in a nursery with three electronic balance with 0.0001 g sensitivity. The color replicates of the dry seeds was determined using the standard morphotypes. The experimental units were kept in a color chart published by the Royal Horticulture weaning shed to reduce the rate of evaporation. The Society (1966). Color was coded by 1 if green and 0 if number of seeds of each morphotype that germinated grey. Seed were subjected to a viability test using the was recorded daily over a 30 days period. At the end flotation method in which those seeds that floated on of this period collar diameter, stem height and water after 24 hours of soaking were considered to be number of leaves were measured weekly on five non-viable and were discarded. seedlings selected at random for each morphotype C. bonduc morphotypes were of identified 30 pots from per the each hierarchical of the four and from each replicate block for 45 days. The seed length, width, thickness, weight and color data were subjected to Ascending Hierarchical The five seedlings measured per morphotype and Classification (AHC) using Ward agglomerative replicate were planted in the field following the same method and Euclidian distance using SAS 9.2 design and measured during 180 days. The collar statistical Mahalanobis diameter, stem height and the number of leaves were distance was used to test the distance between pairs measured monthly on two plant select at random for of the morphotypes. Canonical discriminant analysis each morphotype per replicate block for 180 days. was performed on the morphotypes identified from The germination rate of each morphotype was the AHC in order to validate and test the differences calculated each day over 30 days and the data were between the morphotypes. The assumptions of the used to test the effects of time and morphotype on the canonical discriminant analysis were met (the within- germination of C. bonduc seeds using a mixed model group covariance structure was homogeneous for all ANOVA with repeated measures in SAS software morphotypes and data within the morphotypes had (SAS, 2008). In this model, the factor “block” was software (SAS, 2008). multivariate normal distributions). The morphotypes considered to be random, whereas the factor were also described according to their differences, “morphotype” was considered to be fixed. No data using canonical discriminant axes defined by seed transformation on germination percentages was Padonou et al. Page 88 needed because normality and homoscedasticity 80% of the variations between morphotypes (Fig. 2). assumptions were met using the Ryan-Joiner test of The standardized canonical coefficients and the normality, and the Levene test for homogeneity of correlation coefficients between the two canonical variances (Glèlè Kakaï et al., 2006). The effects of axes and the morphological traits of C. bonduc seed morphotype on seedling and plant growth were (Table 1) indicated that the first axis (Can 1) described assessed using the same statistical methods. 91% of the variation discriminated between morphotypes according to seed color. The second axis Results (Can 2) describing 9% of the variation discriminated Identification of C. bonduc morphotypes between morphotypes according to the length, width, Four morphotypes were identified from the hierarchical classification accounting for 58% of the thickness and weight of the seed. On this axis, heavy seeds were often long, wide and thick. information. The results of canonical discriminant analysis performed on the morphotypes of C. bonduc Seeds from morphotypes 1 and 3 were green, while seeds showed that Mahalanobis distances between seeds from morphotypes 2 and 4 were grey. The pairs of the four identified morphotypes were all heavy, long, wide and thick seeds were from highly significant (p ≤ 0.001). morphotype 3 and 4 while the small seeds were from morphotypes 1 and 2. A more detailed description of Table 1. Standardized canonical coefficients (SCC) each morphotype is provided in Table 2. A deep and correlation coefficients between canonical axes analysis of our data indicated that each morphotype (Can 1, Can 2) and the morphological traits of C. identified, was composed of seeds from the two bonduc seed according to the morphotypes. climate zones. However, seeds of morphotype 1 and 3 Variable Length Width Thickness Weight Colour zSimple Can 1 SCC 0.13 -0.21 -0.09 -0.23 4.50 rz 0.06 -0.09 -0.16 -0.20 0.99 came mainly from the Sudanian zone (56% and 63%, Can 2 SCC 0.79 0.56 0.28 0.34 0.12 rz 0.90 0.89 0.71 0.85 0.07 respectively), while morphotype 2 and 4 seeds were mainly from the Guinean zone (58% and 88% respectively). Morphotype 3 had the longest seeds correlation between each morphological traits and its respective canonical axes (mean length 19.29 mm), while the shortest seed were from morphotype 1 (mean length 17.65 mm). The heaviest seeds were found in morphotype 4 (mean The morphotypes identified were thus highly significantly different according to morphological traits of C. bonduc seed. Other results from canonical discriminant analysis performed on individuals of the four morphotypes revealed that the first two axes were highly significant (P ≤ 0.001) and explained weight 2.85 g), whereas the lightest ones were from morphotype 1 (mean weight 1.95 g). With regard to seed thickness, seed of morphotype 4 were the thickest (mean thickness 15.27 mm), while morphotype 1 seed were the thinnest (mean thickness 13.46 mm). Table 2. Mean values and standard deviations of the morphometric traits of four morphotypes of C. bonduc seed. Traits Morphotype 1 Morphotype 2 Morphotype 3 Morphotype 4 Guinean zone Sudanian zone m s m s m s m s m s m Guinean zone (%) 44 - 58 - 37 - 88 - - - - - Sudanian zone (%) 56 - 42 - 63 - 12 - - - - - Length (mm) 17.7 0.76 17.89 0.75 19.29 0.6 19.28 0.45 18.3 0.92 18.4 1.09 Width (mm) 15.8 0.99 16.52 1.14 18.04 0.7 18.64 0.65 16.9 1.47 16.9 1.32 Thickness (mm) 13.5 0.79 14.05 0.8 14.75 0.9 15.27 0.75 14.2 1 14.1 1.03 Weight (g) 1.95 0.26 2.2 0.33 2.52 0.3 2.85 0.17 2.29 0.45 2.24 0.34 Colour Green Grey Green Grey Grey s Green All values are means +/- SD (n = 2000) Padonou et al. Page 89 Differences between the climate zones according to describing seed morphology according to seed color. Seeds from the Guinean zone The results of canonical discriminant analysis were mostly grey while those from Sudanian zone performed on the seed morphology of C. bonduc were mostly green (Table 2). Seeds length varied from according that 18.38 mm in Sudanian zone to 18.26 mm in Guinean Mahalanobis distance between the two climate zones zone. The mean width of the seeds was 16.91 mm in was highly significant (p ≤ 0.001). Seed morphology Guinean zone and 16.88 mm in Sudanian zone. The of C. bonduc was thus highly significantly different mean tick of seeds from Guinean zone was 14.18 mm according the climate zones. while it was 14.13 mm in Sudanian zone. The weight to the climatic zones showed discriminated between morphotypes of the seeds varied from 2.29 g in Guinean zone to Table 3. Standardized canonical coefficients (SCC) 2.24 g in Sudanian zone. and correlation coefficients between canonical axes (Can 1) and the morphological traits of C. bonduc Germination ability of C. bonduc seeds according to seed according to climate zones. morphotype Variable The germination ability of each morphotype varied in time Can 1 SCC rz Length 0.39 0.24 Width -0.01 -0.03 Thickness 0.13 -0.09 Weight -0.49 -0.25 Colour 0.88 0.94 between sowing and 30 days after sowing (Table 4). Table 4. ANOVA with repeated measures related to the germination ability of the four morphotypes of C. bonduc seed Source DF Time (T) 5 Other results from canonical discriminant analysis Block (B) performed on individuals of the seeds revealed that the first axe was highly significant (P ≤ 0.001) and zSimple correlation between each morphological traits and its respective Type Mean III SS Square 1952.31 390.46 3400.81*** 2 74.23 37.11 45.55ns TxB 10 23.82 2.38 20.75ns Morphotype 3 20.79 6.93 8.51*** TxM 15 22.65 1.51 13.15*** BxM 6 44.28 7.38 9.06ns TxBxM 30 24.77 0.83 7.19*** canonical axes F-value (M) explained 100% of the variations between the climate zones (Fig. 2). The standardized canonical coefficients and the correlation coefficients between the canonical axe and the morphological traits of C. bonduc seed DF, degree of freedom; Type III SS, Type III Sum of Squares; F-value, Fisher (Table 3) indicated that the first axis (Can 1) value; ns, non-significant at P ≥ 0.05; ***, significant at P ≤ 0.001. Table 5. ANOVA with repeated measures related to seedling growth on collar diameters, heights and number of leaves in four morphotypes of C. bonduc seed. Collar diameter Height Number of leaves Source DF Mean Square F-value Mean Square F-value Mean Square F-value Time (T) 8 3.23 58.48*** 278.69 555.15*** 24.00 75.16*** Block (B) 2 2.50 2.14ns 1.38 1.24ns 6.17 0.57ns TxB 16 0.04 0.78ns 0.63 1.27ns 0.31 0.98ns Morphotype (M) 3 2.66 2.27ns 9.41 8.41** 0.38 0.04ns TxM 24 0.06 1.13ns 0.61 1.23ns 0.16 0.51ns BxM 6 0.10 0.09ns 4.30 3.85* 14.16 1.30ns TxBxM 48 0.03 0.60ns 0.63 1.27ns 0.28 0.89ns DF, degree of freedom; F-value, Fisher value, ns, non-significant at P ≥ 0.05; *, significant at P ≤ 0.05; **, significant at P ≤ 0.01; ***, significant at P ≤ 0.001. The blocking factor and the two ways interactions homogeneity of the environmental characteristics with the block were non-significant, indicating between blocks. Germination of the seeds started 5 Padonou et al. Page 90 days after sowing and reached their maximum in 20 (Table 5). The blocking factor and all its interactions days were non-significant indicating homogeneity of the (Fig. 2). The germination rate varied significantly in time, between morphotypes, between environmental morphotypes in time and between block and Morphotypes seedling growth in terms of collar morphotypes in time. The percentage of germination diameter, height and number of leaves varied varied from 80% (morphotype 2 and 4) to 88% between the end of germination (30 days after (morphotype 1) after 30 days. sowing) and 45 days later (Fig. 4). C. bonduc seedling growth according to morphotype Plant growth of C. bonduc according to the No difference was observed between morphotypes in morphotypes number of leaves and collar diameter, while the There are significant differences in plant growth growth in height differed between morphotypes between morphtypes (Table 6). characteristics between blocks. Table 6. ANOVA with repeated measures related to plant growth on collar diameters, stem heights and number of leaves in the four morphotypes of C. bonduc seed. Collar diameter Height Mean Mean F-value Number of leaves Source DF Time (T) 5 156.63 42.28*** 2504.47 30.68*** 343.61 18.78*** Block (B) 2 6.38 12.19* 20.04 4.83* 9.36 0.62ns TxB 10 0.70 0.19ns 25.46 0.31ns 10.30 0.56ns Morphotype (M) 3 22.52 42.98*** 1124.23 24.70*** 34.35 8.34** TxM 15 1.74 0.47ns 197.55 2.42** 34.35 1.88* BxM 6 1.95 3.74ns 112.18 2.46ns 50.97 3.40* TxBxM 30 0.70 0.19ns 64.95 0.80ns 14.89 0.81ns Square F-value Square Mean F-value Square DF, degree of freedom; F-value, Fisher value; ns: non-significant at P ≥ 0.05; *, significant at P ≤ 0.05; **, significant at P ≤ 0.01; ***, significant at P ≤ 0.001. Plants of morphotype 3 have the highest values of indicating collar diameter and height. The highest values of the characteristics between blocks. Growth in terms of number of leaves were observed with morphotype 4, collar diameter, height and number of leaves varied followed by morphtype 3 (Figure 5). The blocking between the end of seedling period and six months factor and all its interactions were non-significant, later for all four morphotypes (Figure 5). homogeneity of the environmental Figure 2. Projection of the four morphotypes of C. bonduc seed on the canonical Figure 3. Cumulative germination rates of C. bonduc seed axes defined by seed morphology. Can 1 discriminated between morphotypes according to morphotype. according to seed color. Can 2 discriminated between morphotypes according to the length, width, thickness and weight of the seed Padonou et al. Page 91 Figure 4. Trends in seedling traits over time (in days) following Figure 5. Trends in plant traits over time (in days) following germination in C. bonduc based on collar diameter (Panel a), height (Panel seedling growth in C. bonduc based on collar diameter (Panel a), b) and leaves (Panel c) according to seed morphotype. height (Panel b) and leaves (Panel c) according to seed morphotype. Discussion grey and presented more compact morphological Morphological variation in C. bonduc seed traits (shortest, thickest, widest, and heaviest) than C. concerning those from Sudanian climatic zone. However, none of morphological seed traits. Four morphotypes of seed the four morphotypes had characteristics which fitted were significantly distinct according to some of their perfectly those of Guinean or Sudanian seed traits. morphological traits. Seed of morphotypes 3 and 4 Thus the overall environmental factors cannot explain were heaviest, longest, widest and thickest, in the substantial morphological variation found within contrast to morphotypes 1 and 2 which were smaller. populations (Assogbadjo et al., 2012). These results This canonical are consistent with a recent study in Benin on the discriminate analysis confirmed the statement that morphological and genetic diversity of C. bonduc tropical bonduc showed differentiation variation confirmed with important trees, which reported that the species is characterized intraspecific variation of seed traits (Foster, 1986; by a high morphological variation among individuals Khan et al., 1999, 2002; Khan and Uma, 2001; Khan, of the same populations, while a much lower degree 2004). The results corroborated with recent studies of on other species such as Jatropha curcas (Ginwal et populations and climatic zones (Assogbadjo et al., al., 2005; Padonou et al., 2014), Afzelia africana 2012). According to Mathur et al. (1984), the plant al., often have variation were observed among the indica morphological difference between seeds could be of genetic origin, due to the adaptive strategies of 2007). Indeed, seeds from Guinean zone were mostly species to their environment. But Assogbadjo et al. 2013) Tamarindus such (Fandohan, 2010) and Prunus jenkinsii (Upadhaya, (Padonou et species Padonou et al. Page 92 (2012) revealed that C. Bonduc is characterized by a greatest collar diameter, height and number of leaves very low genetic diversity within populations. The of plant. We could suggest that seed mass could phenotypic plasticity of the species in response to explain observed performance of morphotype 3. micro habitat factors, and the difference of age of Many authors reported that heavier seeds possess targeted observed large reserves of nutritive substances available, which differences of seed morphology (Heaton et al., 1999; confer an advantage to their seedlings for survival and Assogbadjo et al., 2005; Assogbadjo et al., 2012). growth (Tripathi and Khan, 1990; Ke and Werger, individuals can explain the 1999; Khan et al., 1999, 2002; Khan and Uma, 2001). Seed germination, seedling and plant growth in C. This advantage is a good start which remains during bonduc seed plant growth phase. Nevertheless bad performance of This difference in germinataion rate could be seeds of morphotype 4, which were heavy seeds too, explained by morphology, which is an important supposed that good germination percentage of C. factor influencing the germination rate. Many reports bonduc seeds, influenced juvenile and plant growth existed on the conflicting relationships between the phase later. variation in seed weight and germination behaviour. For instance, large seeds may germinate at higher Conclusion percentages than small seeds (Tripathi and Khan, The study revealed that considerable variation exists 1990; Bhuyan and Khan, 2000; Khan and Uma, in C. bonduc populations with respect to seed 2001), and small seeds may germinate at higher morphology. Except seed color which varied from percentages than large seeds (Marshell, 1986), or grey in Guinean zone to green in Sudanian zone, there germination may be independent of seed size (Gross were no significant differences between the two and Kromer, 1986; Perez-Garcia et al., 1995). Since climatic zones on the others traits. Seed germination morphotype 1 is the lightest and morphotype 4 is the was not significantly related to seed morphology. heaviest, we could conclude that our data revealed an Nevertheless, independent relationship between germination and influenced by seed morphology. Moprhotype 3, which seed which consists of large green seeds mostly from the discriminated these two groups, and could explain the Sudanian zone is superior in terms of seedling and difference observed, is the color of the coat. Indeed plant and should be the best choice for planting morphotypes 1 and 3 were green while the two others purposes. mass. Beside the main factor seedling and plant growth were had a grey coat. Similar results were obtained by Mavi (2010) who revealed that the seed coat color of References Crimson sweet seeds could explain the differences of Adomou seed quality. Moreover, the grey coat color was Environmental gradients in Benin. Implications for mainly characteristic of seeds from Guinean climatic biogeography zone while green coat indicated seeds collected from Wageningen University, Wageningen, p. 150. CA. 2005. Vegetation Patterns and and conservation. Ph.D. Thesis, Sudanian. But these hypotheses need additional research to test. On one hand, for Crimson sweet Anonymous 1956. Wealth of India Delhi. Council of seeds, the seeds color (brown) was coupled with high Scientific and Industrial Research Publications, p. 6- seeds mass, to explain good germination rate. But in 8. our case, seed size do not influence germination percentage. Assogbadjo AE, Fandohan B, Glèlè Kakaï R, As far as seedling and plant growth concern, seeds of Kyndt T, Hardy OJ, Gheysen G, Sinsin B. 2012. morphotype 3 confirmed their vigour and quality. 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