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Cuban Journal of Agricultural Science

Print version ISSN 0864-0408On-line version ISSN 2079-3480

Cuban J. Agric. Sci. vol.55 no.4 Mayabeque Oct.-Dec. 2021  Epub Dec 01, 2021


Pasture Science and other Crops

Characterization of the floral structure and evaluation of seeds production from different materials of Tithonia diversifolia Hemsl.) A. Gray. in Cuba

Idalmis Rodríguez García1  *

C. Padilla Corralea1

Yolaine Medina Mesa1

1Instituto de Ciencia Animal, Apartado Postal 24, San José de las Lajas. Mayabeque, Cuba


In order to characterize the reproductive structure, the production and seed germination of different materials of Tithonia diversifolia Hemsl.) A. Gray, a total of three experiments were performed at different times and five plant materials were evaluated (3, 10, 16, 23, 25). The harvest of ten heads (2018 and 2019) was carried out by means of a random sampling at five points on the diagonal of the field. The evaluated materials constitute the treatments. A completely random design was used for the analysis of the variables diameter (cm) and seed heads weight (g), number and weight (mg) of full and empty seeds and of PGS seed heads-1. To the total number and the germination percentage of seed at different times 3, 5, 7 and 30 days) a simple classification non-parametric analysis of variance was performed. The studied materials showed phenotypic differences in terms of their reproductive structure. The seed heads of material 25 had a higher diameter. Material 16, due to its higher weight, distinguished itself from the rest of the evaluated materials. The waste from the seed heads is approximately 71.18 % of their total weight. The seed production of materials 10, 23 and 25 is very similar with respect to the total number of seeds in the seed heads, as well as the number of empty seeds. However, material 23 showed a lower number of full seeds (P ≤ 0.0005). These results contribute to the knowledge of the reproductive characteristics of each specific material, which will allow the development of future studies related to the gamic seed production strategy.

Key words: gamic seed; Tithonia; germination; inflorescences

In Cuba, Ruíz et al. (2017) determined the characteristics that are contrasting, in terms of morphological traits and seed germination percentage, from a group of Tithonia materials, collected in different edaphoclimatic regions of the country (29 of the central-western zone in 2006 and 52 of the central-eastern zone in 2015).

In the world today, this plant is of great interest for animal feeding. In Cuba, it is considered one of the cheapest ways to supply protein, so the search for economic alternatives for the reproduction and propagation of this species through gamic way constitutes a challenge and a priority for the scientific community.

The objective of this research was to characterize the reproductive structure, production and seed germination from different materials of Tithonia diversifolia Hemsl. A. Gray.

Materials and Methods

The research was developed in a red eutric ferralitic soil (Hernández et al. 2015), at the Miguel Sistach Naya experimental station, from Instituto de Ciencia Animal, located in Mayabeque, Cuba. The plant materials of T. diversifolia corresponded to those proposed by Ruíz et al. (2014) for cut or grazing. The characterization of the floral structure was determined in three researchers and the evaluation of seed production was developed in experiment 2.

Experiment 1. The samplings were carried out in plots of 0.028 ha, established in August 2018, with the objective of producing seeds. The evaluated materials were 10, 23 and 25. They were harvested in an area of 50 seed heads-1 of the flowering of November 2018.

Experiment 2. In the same field of the previous experiment, a total of 50 heads-1 from November 2019 flowering were harvested.

Experiment 3. The materials evaluated were 3, 10, 16 and 23. The inflorescences were collected from fields of T. diversifolia, established in July 2015. The sampling area consisted of five 25 m2 plots.

In experiments 2 and 3, the harvest was carried out in previously established fields. For this, a uniformity cut of the area was made in May, with the intention of causing the appearance of uniform flowering flows, which occurred in the last week of October in all the studied materials. The best time of seed harvest was reached in the first half of November.

The sampling was carried out according to the phenological state of seed heads, proposed by Padilla et al. (2018) (green seed heads with withered petals and without petals and seed heads with bracts and dry peduncles, brown). A total of 10 seed heads were taken at five sampling points, randomly and diagonally. The diameter (cm) was determined with a vernier, as well as their green weight (g). Then, the drying and postharvesting was carried out to obtain the seeds. From 10 heads, the DM percentage, total dry weight (g), waste weight (difference between the total weight of heads and the seeds weight, g), weight of full and empty seeds (mg) and of the PGS seed heads-1 were evaluated. The latter were calculated from the weight of full seeds plots-1 and the germination percentage and the weight of 1000 full seeds (g). The weight of seeds and seed heads was determined on an analytical balance (TE12S, Sartorius).

The total number of seeds was quantified and they were classified as full and empty, determining their hardness to the touch. The germination percentage was evaluated by means of a germination test, placing 100 full seeds per Petri dish and four replications, as indicated by the International Seed Testing Association standard (ISTA 2015). A cotton substrate was used which was moistened with distilled water. The dishes were placed under laboratory conditions, at room temperature (26-28 ºC) and natural light. The percentage of total germination at 30 days (and at three times at 3, 5 and 7 days after germination), the number of germinated seedlings in the dishes and the DM percent were determined.

The theoretical assumptions of the analysis of variance were verified for all variables, from Shapiro and Wilk (1965) test for the normality of the errors and the Levene (1960) test for the homogeneity of variance.

The variables diameter (cm) and total weight of seed heads (g), number of full and empty seeds, the weight of each one and of the PGS seed heads-1 fulfilled the assumptions. The ANAVA was performed according to a completely random design, where the evaluated materials constituted the treatments. Duncan (1955) test was used to compare the means. The data were processed using the statistical package Infostat (Di Rienzo et al. 2012).

The variables total number and germination percentage of seeds (at different times and total) did not fulfill with the theoretical assumptions of the ANAVA. Subsequently, the data transformations √x were applied for the variable total number of seeds and arcsin √%, for the variables expressed in %. However, these ones did not improve this compliance, so a simple classification non-parametric analysis of variance was performed (Kruskal Wallis) and the Conover (1999) test was applied to compare the mean ranges.

The variables number of seedlings and their DM percent did not fulfill with the theoretical assumptions of the ANAVA. The √x and arcsin √% transformation did not improve the cited assumptions, so a non-parametric random block analysis of variance (Friedman) was performed, with a 3*3 factorial arrangement. The factors were the evaluated materials (10, 23 and 25) and the moments the number of germinated days (3, 7 and 10). For the comparison of mean ranges, the Bonferroni (1936) test was applied.

Results and Discussion

Little is known about the floral structure of the capitulum or seed head, the anatomy of flowers and the developmental changes of the floral organs of T. diversifolia. However, researchers of other Asteracea species, performed by Oba et al. (2017) and Ibañez et al. (2017) on the external and internal structure of flowers and the characterization of the development stages of the seed heads allowed to better understand the reproductive characteristics of this family, and to clarify the cause of the low efficiency of seed propagation. In Cuba, Padilla et al. (2018) showed in T. diversifolia that from the phenological changes that occur in the heads, the optimal harvest time can be defined, and with this it is possible to significantly increase the germination percentage, weight of 1000 seeds and yield of PGS. The mentioned authors, based on their results, clarified the prevailing criterion, which states that the seeds of this species have low germination and viability.

The studied materials have phenotypic differences in terms of their reproductive structure (table 1). The heads from material 25 showed higher diameter and green weight of the heads (table 2), with respect to the other materials. However, they did not differ from material 23, in terms of weight (GM and DM) and DM percentage of the floral structure (table 2 and 3). Material 16 showed phenotypic characteristics, in terms of its reproductive structure (diameter and weight of the seed heads), which distinguished it from the other materials evaluated in experiment 3.

Table 1 Performance of seed heads diameter (cm) of different materials 

Experiments Materials SE± p
10 23 25 3 16
1 1.95c 2.05b 2.36a - - 0.019 <0.0001
2 2.43b 2.35c 2.97a - - 0.027 <0.0001
3 2.45b 2.34c - 2.51b 2.58a 0.022 <0.0001

Means with different letters in each row differ at P ≤ 0.05 Duncan (1955)

Table 2 Green weight of seed heads from different materials, g 

Experiments Plant materials SE± p
10 23 25 3 16
1 0.18 0.15 0.19 - - 0.01 0.13
2 0.11b 0.10b 0.42a 0.03 <0.0001
3 0.31b 0.30b - 0.32b 0.44 a 0.02 0.0001

Means with different letters in each row differ at P ≤ 0.05 Duncan (1955)

Table 3 Weight and DM percentage of floral structure from three Tithonia materials 

Variables Materials SE± p
10 23 25
Weight GM heads, g 1.88 1.50 1.95 0.07 0.07
Weight DM heads, g 0.59b 0.64ab 0.75a 0.01 0.02
Waste weight (DM) heads, mg 0.42b 0.51a 0.53a 0.01 0.02
DM percentage heads, % 34.16b 44.11a 39.94a 0.04 0.07

Means with different letters in each row differ at P ≤ 0.05 Duncan (1955)

According to González-Castillo et al. (2014), the heads can reach up to four cm in diameter. In this case, the values showed in the literature are between 2.5 and 3.5 cm. Dispersion studies of this species, developed by Wang et al. (2004 and 2008) in five geographically different regions of China, point out that the diameter of the seed heads was 2.6 to 3.2 cm.

The waste weight (table 3) of heads is approximately 71.18 % of the total weight. It is very important to know this data for the processing of seeds, since the amount of seeds wastes to be separated during the benefit process is considerable. Materials 23 and 25 showed a higher DM percentage and a higher weight of waste material, which could be associated with the specific characteristics of the material.

The high DM content in the seed heads is an important trait to consider. According to Marcos Filho (2015), this variable can influence on the process of seed formation, considering that a higher accumulated dry mass is a signal that the seeds have reached their physiological maturity, when there is no more translocation of assimilates from the plant to the seeds. In this sense, Morales-Nieto et al. (2012) show that the seed weight is the best indicator to define that they achieved their physiological maturity and are suitable for harvest.

According to Romero-Saritama (2018), morphological and physiological studies in seeds from woody species are of great importance and their knowledge allows relating the adaptation of these species to the edaphoclimatic conditions where they develop and, in turn, makes it possible to draw strategies for its conservation and extension. This author studied the regenerative morphological traits of seeds from different species in the dry forests of Ecuador. Among the main traits or characteristics that he evaluated are the tolerance to desiccation (most of the species have low moisture content, between 3 and 7 %), the size (average between 10 x 6 mm in length and width respectively), weight (mass less than 3 g) and shape (presence of developed embryos).

The morphological traits of Tithonia seeds, like those of most Asteraceae, allow a greater dissemination of them. According to Santos-Gally et al. (2020), heteromorphism (differences between central and peripheral achenes in a head) in T. diversifolia could represent a strategy of this species for its adaptation to uncertain environmental conditions. These authors point out that the central achenes have beard (potentially associated with zoocorous dispersion), weak latency and higher germination rate and success, at least in one of the two evaluated populations. Peripheral achenes lack structures associated with dispersion and have a lower germination and success rate.

Studies carried out in Rwanda (Seburanga 2014) showed that up to 18 % of the immature achenes of a head can be removed by canaries (Serinus capistratus), so it is necessary to consider the affectations that can be presented by birds in seed production.

In the three materials (figure 1), seeds production was very similar with respect to the total number of seeds in the heads, as well as in relation to the number of empty seeds. However, material 23 showed a lower number of full seeds (P ≤ 0.0005).

Figure 1 Gamic seed production of three Tithonia materials 

In each of the evaluated materials, the number of full seeds, determined by inflorescence (head), was below that reported in the literature for other regions. In China, Wang et al. (2004) determined between 164.2 and 231 seeds per heads. In Nigeria, Muoghalu and Chuba (2005) and Muoghalu (2008) showed values of 179.7 and between 136 - 144 seeds, respectively. According to Mattar et al. (2019), each inflorescence of T. diversifolia can produce about 189.8 seeds.

In other researchers, values closer to the range determined in this study are reported, such is the case of the studies of Etejere and Olayinka (2014 and 2015) (32-62 seeds seed heads-1) in Nigeria and Silva et al. (1990) (89.9 seed heads-1) in Brazil. In Cuba, studies by Padilla et al. (2018), where the optimal harvest time is defined according to the phenological development of the seed head maturity, the number of full seeds per seed head was 11, 31 and 23, when they were harvested in the state of seed heads with green bracts and withered petals, green bracts without petals, and dry brown bracts and peduncles, respectively. It was showed that in the crops in the youngest stages of seed head development, the presence of full seeds was lower (11) with respect to that of empty seeds, which was higher (53). It was also observed that in the more mature seed heads it is reduced to 22 empty seeds per seed head.

However, the above, in dispersion studies of the populations of this species (Sun et al. 2007), carried out in China, higher values of seed production were reported (80 000 to 160 000 achenes m-2). According to Gallego-Castro (2016), in the edaphoclimatic conditions of the high tropics from Antioquia region in Colombia, the average number of flowers per plant is 88.7, and each flower produces, on average, 61.2 seeds. The discussion of the indicators number of seed, germination percent and weight of heads, as well as the weight of 1000 seeds depends on the type of seed and the precise information (full or total seeds and the purity of the lot), which sometimes it is not showed in the researchers methodology.

In Colombia, Saavedra (2016) characterized the germination performance of the sexual seed of T. diversifolia from different sources. To do this, he collected seeds in four departments, where the ancient presence of the species has been recorded (Meta, Antioquia, Caldas and Valle del Cauca). The seed viability tests showed that more than 15 % of the produced fruits are empty, more than 9 % of the seed is rudimentary, and more than 50 % of the produced seeds have some malformation that makes them non-viable. Therefore, approximately 74 % of the fruits produced by this species do not form a seed, and do not develop into new individuals.

The material 25 highlighted for the higher weight of 1000 seeds (6.54 g) (table 4). The values of the weight of 1000 seeds were higher than those determined by Ayeni et al. (1997). According to these authors, the populations of T. diversifolia that grows in the south west of Nigeria have values between 2.5 and 2.6 g. Wang et al. (2004) and Sun et al. (2007), in China, determined figures between 4.64 and 6.50 g, when carrying out studies in various regions of Yunan province. Gallego (2016) reports a weight of 7.36 g per 1000 seeds, which is higher than that obtained in this research. The materials 10 and 25 do not differ in terms of the weight of PGS per seed heads, which may be associated with the similar weight obtained from the full seeds.

Table 4 Weight of 1000 seeds and PGS heads-1 of three Tithonia materials 

Variables Materials SE ± p
10 23 25
Weight of 1000 seeds, g 5.26b 5.69b 6.54a 0.20 0.0005
PGS, mg per heads 3.79a 1.60b 5.48a 1.13 0.0014
Weight of full per heads, mg 0.09a 0.03b 0.09a 0.008 0.0010
Weight of empty seeds per heads, mg 0.07b 0.08ab 0.11a 0.007 0.0400

Means with different letters in each row differ at P ≤ 0.05 Duncan (1955)

In Cuba, Ruíz et al. (2018) report on the germination capacity of some of the studied materials. Regarding the morphological characteristics and seed germination, Rivera et al. (2018) determined for the material 25 values higher than those of this study (73%) (table 5). However, for materials 10 and 23, the germination percentages obtained here were higher (44 and 53 % respectively) than those reported by these authors (6.5 and 20 %).

Table 5 Germination percentage of different T. diversifolia materials 

Indicator Plant material p
10 23 25
Percentage of total germination

  • 10.60

  • (44.22)

  • SD =8.15

  • 16.20

  • (53.61)

  • SD =10.97

  • 19.70

  • (60.90)

  • SD =20.31

Germination percentage 72hours

  • 16.20ab

  • (37.10)

  • SD =6.08

  • 10.15b

  • (27.50)

  • SD =10.54

  • 20.15a

  • (51.10)

  • SD =23.38


Means with different letters in each row differ at P ≤ 0.05 (Bonferroni 1936).

SD= Standard deviation ( ) mean ranges

In Mexico, Santos-Gally et al. (2017, 2019) obtained in this species germination percentage higher than 90 % in laboratory germination tests, while in field sowings they were between 50 and 70 %. A higher germination percentage (81 %) reached Etejere et al. (2014), when sowing sexual seed in pots. These authors determined that the sowing depth is a factor that significantly affects the germination percentage. Likewise, they evaluated the germination response at different depths (0, 0.5, 1.5, 2.5 and 3.5 cm) with respect to the soil surface. They observed that the highest germination percentage occurred at depth 0 cm, and decreased as depth increased. At depths of 0 to 2.5 cm, the emergence occurred approximately five days later, and at depths of 3.5 cm it started from the sixth day.

Padilla et al. (2020) propose to cover the seed with small amounts of cattle manure in each niche, to promote stable conditions for the germination and survival of young plants, which causes a better development of them.

The non-parametric analysis of variance of random blocks with factorial arrangement did not show significant interaction, so the main effects for the variable number of seedlings and DM percentage can be seen (table 6).

Table 6 Number of germinated plants and DM percentage of the seedlings from different T. diversifolia materials 

Indicators Materials Moments: number of germinated days
10 23 25 3 5 7
Number of seedlings

  • 30.60c

  • (5.50)

  • SD=24.49

  • 43.67b

  • (7.93)

  • SD=18.76

  • 60.68a

  • (16.67)

  • SD=24.57

  • 54.82a

  • (15.70)

  • SD=29.92

  • 44.75ab

  • (8.87)

  • SD=23.42

  • 35.10b

  • (5.66)

  • SD=19.66

p <0.0001 0.0006
% DM

  • 32.36b

  • (4.14)

  • SD=5.23

  • 59.67a

  • (10.94)

  • SD=6.53

  • 45.37ab

  • (8.05)

  • SD=3.08

  • 45.33

  • (8.60)

  • SD=6.81

  • 40.89

  • (6.69)

  • SD=5.70

  • 50.89

  • (7.73)

  • SD=4.66

p <0.0001 0.2246

Means with different letters in each row differ at P ≤ 0.05 (Bonferroni 1936).

SD= Standard deviation ( ) mean ranges

Seedling emergence is probably the most important phenological event that influences on the success of a plantation. It represents the moment in which a seedling becomes independent of the non-renewable seminal reserves, originally produced by its parents, and it is when photosynthetic autotrophism begins.

Once the emergence of the radicle has occurred and the seedling growth has started, the latter uses the nutrient reserves stored in the seed during the development phase, in order to support its growth. The efficiency with which this process occurs is probably related to the vigor and growth rate of the seedling, which in turn influences on the probability of a successful emergence in the field and in the establishment of the plant (Hilhorst and Bradford, 2000).

In material 23, despite having the lower number of seeds per seed heads and weight of PGS, it can be seen that the DM percentage of seedlings was higher. Similarly, the seedlings from material 25 showed higher values. This performance and its relation with the vigor and growth rate of the seedlings can be associated with a physiological response of the material to guarantee greater survival.

It is concluded that the endorsed materials show differences, regarding the floral structure and the number of seeds per seed heads, as morphological characteristics, and in their germination.

The results obtained in this study contribute to the knowledge of the reproductive characteristics of this plant and of each specific material, which will allow the development of future studies related to the gamic seed production strategy.


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Received: June 25, 2021; Accepted: October 04, 2021


Conflict of interest: The authors declare that there are no conflicts of interests among them

Author´s contribution: Idalmis Rodríguez García: Original idea, conducting the experiment, writing the manuscript. César Padilla Corrales: Experimental design, writing the manuscript. Yolaine Medina Mesa: Statistical analysis

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