Temperatures and light regimes in the germination of Areca vestiaria and Areca triandra seeds¹

ABSTRACT

Propagation of palm trees is mainly by seeds and, in general, germination is uneven and slow, caused by several factors, such as temperature and light. We aimed to evaluate the effect of different temperature conditions and light regimes in the germination of Areca vestiaria and Areca triandra seeds. Two experiments were carried. The experimental design was completely randomized with a 5 × 2 factorial (temperature × light), with four replicates per treatment for the species A. vestiaria and five replications for A. triandra, with 25 e 20 seeds, respectively. The temperatures tested were 25, 30, 35, 20-30 and 25-35 ºC and the light regimes were presence and absence of light. The germination percentage and germination speed index were evaluated. Analysis of variance was performed using the F Test, with subsequent comparison of the means using the Tukey test (α = 0.05). For both species, there were significant differences between the temperature conditions for the studied characteristics. It was concluded that temperatures of 25 ºC, 30 ºC, 25-35 ºC and 20-30 ºC are indicated for germination of seeds of the species Areca vestiaria and, 25-35 ºC for Areca triandra, being the seeds classified as neutral photoblastics.

Keywords
Arecaceae; ornamental plants; photoblastia; palm trees; thermal stress

INTRODUCTION

Palm trees have great variability in species richness, phylogenetic composition and life forms because they are adapted to a variety of climates and soils (Eiserhardt et al., 2011). The Arecaceae include approximately 2700 species in 240 genera (Lorenzi et al., 2010). Due to its exuberance, beauty, size and crown that stand out in nature, they have great ornamental potential, providing remarkable harmony in the landscape compositions. Due to these characteristics, all palm trees are considered ornamental, although some are widely used and others are unknown (Costa et al., 2018).

Areca vestiaria Giseke, originally from eastern Indonesia, is a cespitose palm tree, occasionally solitary, monoecious, forming thin or compact clumps and quite attractive due to the unique color of the leaf sheath in the palm heart region. Areca triandra Roxb. ex Buch-Ham., originally from Southeast Asia, East India, Sumatra, Borneo and the Philippines, is a cespitose palm with detached nodes and internodes similar to bamboo, sometimes with adventitious roots at the base, as well as flowers with a characteristic scent of lemon (Lorenzi et al., 2004).

The propagation of palm trees is mainly by seeds and in general they have uneven and slow germination, caused by several factors (Meerow & Broschat, 2015). Temperature, humidity, light and oxygen are considered fundamental for its germination (Carvalho & Nakagawa, 2012). For Peske et al. (2012), the ability of seeds to germinate under a wide range of environmental conditions ensures the survival and regeneration of species.

Germination is dependent on well-defined temperature limits, characteristic for each species (Bewley et al., 2013), considering that, at optimal temperature, the percentage of germination is higher and, in less time, already at maximum and minimal temperatures, there may be a lower percentage of germination or death of the embryo (Carvalho & Nakagawa, 2012). Temperature interferes in the dynamics of water absorption, as well as in the limits and speed of biochemical reactions and, also, in the physiological processes that determine the germination of seeds (Marcos Filho, 2015).

Regarding the light stimulus in the germination process, the seeds that need light are called positive photoblastics, while those that germinate better in the absence of light are negative photoblastics and, when there is no light interference in the germination, the seeds are considered neutral photoblastics (Nogueira et al., 2014). The light participates in the activation of the seeds and is linked to phytochrome, which is the pigment responsible for capturing the light signals from the environment (Brancalion et al., 2012), which can have a stimulating or inhibiting effect on germination, depending on the light wavelength at which it was submitted (Carvalho & Nakagawa, 2012).

Temperature and light conditions can be controlled and, thus, to contribute to improving the percentage, speed and synchronization of germination. In view of the need to determine which conditions are most suitable for seed germination for the establishment of a reliable protocol, especially for species of the Arecaceae Family.

The objective of this study was to evaluate the effect of different temperature conditions and light regimes on the germination of Areca vestiaria and Areca triandra seeds.

MATERIAL AND METHODS

The fruits of Areca vestiaria were harvested from parent plants grown on a property located in Brumadinho city, Minas Gerais State (20º7’26.86” S and 44º13’7.63” O), on October 13th, 2019 and from the species Areca triandra from plants grown at the Experimental Nursery of Ornamental and Forest Plants at São Paulo State University (Unesp) (21°15’2’’ S and 48°16’47’’ O), Universidade Estadual Paulista, Jaboticabal, São Paulo State, on February 12th, 2020. Six plants matrices were randomly selected for each species. The fruits of both species were harvested ripe, evidenced by the red color of the epicarp and loosening of the bunches (Lorenzi et al., 2004).

Two experiments were carried out at the Laboratory of Horticultural Seeds, located in the Department of Agricultural Sciences, Plant Production Sector, FCAV/UNESP, Jaboticabal, SP. The experiments were carried out separately for each species, at different times. In the laboratory, the fruits were pulped (removal of the epicarp and mesocarp) through manual friction with a steel mesh sieve (6 mm) (Beckmann-Cavalcante et al., 2012). Fruit depulping was done three days after harvest for A. vestiaria and, for A. triandra, seed extraction was done on the day of harvest. For both, sowing was carried out on the day after depulping. The diaspores subsequently underwent asepsis by soaking in sodium hypochlorite solution (2%) for 10 minutes, then they were rinsed with running water. This process aims to reduce fungal infestation in seeds (José et al., 2012).

For both, the experimental design was completely randomized with a 5 × 2 factorial, with four replicates per treatment for the species A. vestiaria and five for A. triandra, with 25 and 20 seeds, respectively, totaling 1000 seeds for each species. The combinations of temperatures were 25, 30, 35, 20-30 and 25-35 ºC and two light regimes, it being in the presence: photoperiod with a regime of 8 hours with light and 16 hours without light; and total absence of light. At alternate temperatures, those higher (30 °C and 35 °C) corresponded to the daytime temperature, while the lowest (20 °C and 25 °C) to the night. The number of repetitions and seeds in each treatment differed between species because A. triandra has larger seeds than A. vestiaria, thus needing more repetitions.

Transparent and black plastic boxes with lids (11 × 11 × 3 cm) were used for the treatments of presence and absence of light. In the presence of light, the lamps were turned ond and off automatically for a photoperiod of 8 hours of light being in accordance with what was proposed by Luz et al. (2014). The substrate medium-sized expanded vermiculite (Batista et al., 2016; Costa et al., 2018; Almeida et al., 2018), maintained at 100% of its water-holding capacity. The substrate was slowly saturated with water, until the interruption of its drainage was observed to determine the of water retention. The boxes were wrapped in transparent plastic bags and placed in a germination chamber of the Biochemical Oxygen Demand (B.O.D), with their respective temperature and photoperiod conditions for each treatment. The germination chambers used had 4 daylight white fluorescent lamps with 20 watts each (ELETROLAB^®, model EL202/4).

The water content, germination and germination speed index were evaluated. The germination evaluation was performed daily, considering that the seeds that emitted the germinative bud were considered germinated, until germination stabilization for all treatments. The stabilization of the germination process was observed at 52^nd days for A. vestiaria and at 105^th days for A. triandra. For the treatment with total absence of light, the evaluations were carried out in a darkroom, using a flashlight (20 watts) covered with two sheets of green cellophane paper (Coelho et al., 2012). In this case, the light was turned on only at the time of evaluation of the experiments. In order to maintain the randomness of the experiments, at each evaluation the positions of the boxes were changed at random.

To determine the water content of the seeds, for both experiments, two subsamples with 10 seeds each were separated, using the greenhouse method at 105 ± 3 ºC, for 24 hours (Brasil, 2009). This method consists of extracting water from the seeds in the form of steam by applying heat under controlled conditions, and the results are expressed as a percentage. The water contents found in the seeds were 43.09% and 41.01% respectively, for A. vestiaria and A. triandra. To evaluate the Germination Speed Index (GSI), the formula established by Maguire (1962) was used, based on the values ​​of seed count germinated daily, where , where GSI = Germination Speed Index; G1, G2 and Gn = number of seeds germinated in the first, second and last counts; N1, N2 and Nn = number of days elapsed after sowing in the first, second and last count. With tThe percentage of germination percentage was evaluated at the end of the experiments, using the formula proposed in the Rules Manual for Seed Analysis (Brasil, 2009), considering the formula , where %G = germination percentage; N = number of germinated seeds; A = total number of seeds placed to germinate. Results are expressed as a percentage.

The data were submitted to Shapiro-Wilk normality tests and Levene homogeneity of variances (p > 0.05), with the germination percentage values ​​previously transformed into arc sine (x/100)^1/2, due to the failure to meet the assumptions for the variable. Then, the analysis of variance (ANOVA) was performed by the F Test (p ≤ 0.05) and the comparison of the averages by the Tukey test at the significance level of 5% (α = p ≤ 0.05), using the statistical software AgroEstat^® version 1.1. 0.711 (Barbosa & Maldonado Júnior, 2015). Scatter plots were built with the aid of Microsoft Excel^® software version 2016 for better visualization of germination over time.

RESULTS AND DISCUSSION

The interaction between temperature and light was not significant for evaluated with the two species (Table 1). For A. vestiaria, the temperature had significative effect for all characteristics evaluated, with higher germination percentages in the temperatures of 25; 30; 20-30 and 25-35 ºC, when compared to 35 ºC. For the GSI characteristic, it is observed that the temperature of 35 ºC differed statistically only from the temperatures of 30 ºC and 20-30 ºC. It is possible to notice that the germinative process of the seeds submitted to the presence of light was similar to those kept in the continuous dark, not differing statistically its averages for percentage of germination and GSI (Table 1).

It was also observed (Table 1) that the germination percentages are low (below 40%). Several factors may have influenced the results, for instance, the time from harvest to sowing, the form of storage and transport that differed between species, resulting in a possible loss of viability for A. vestiaria seeds. According to the literature for palm trees, in general, sowing should be done soon after the fruit is harvested. Furthermore, the differences in genotype, origin and genetic load of the matrices during fruit filling and the water replacement rate, which may have been different in the studies, may have a direct influence on the physiological quality and viability of the seeds, as reported by Beckmann-Cavalcante et al. (2012). However, for the species of this research, studies on the physiological maturity of its seeds were not found in the literature.

Although it has a range of temperature for germination, it was found that the constant of 35 ºC negatively affected the germination process, as it promoted a lower value for the percentage of germination and GSI for A. vestiaria (Table 1). Marcos Filho (2015) reports that high temperatures can reduce the percentage of germination, and the number of seeds that manage to germinate drops quickly, basically as a result of the effects on enzyme activity and restrictions on oxygen access. In general, the seeds show variable behavior and there is no optimum and uniform temperature for the germination of all species (Bewley et al., 2013), as observed for both species in this study, which demonstrated a wide variation in thermal need for seed germination. Palm seeds generally require high temperatures for fast and uniform germination, considering temperatures in the range from 21 to 38 °C as acceptable and 29 to 35 °C as more favorable (Meerow & Broschat, 2015).

Regarding the species A. triandra, for germination percentage, there was an effect only for the temperature conditions tested (Table 1). It is also noteworthy that there was no germination at a temperature of 25 ºC in both assigned light regimes (Table 1), which demonstrates the sensitivity of the seeds to these conditions, and can be considered as an adaptive characteristic capable of regulating germination in non-ideal times, even being designated as critical for this species. Low temperatures compromise essential pathways for germination, so that the reorganization of the cell membrane system can become slower, influencing water absorption and, consequently, reducing the biochemical and physiological reactions that determine the vigor and germination of seeds (Flores et al., 2014; Carvalho & Nakagawa, 2012).

As in the present work, other palm species have the light factor as a non-limiting factor for the germination of their seeds, thus the seeds are indifferent to light in their germination process, namely, Copernicia alba Morong ex Morong & Britton (Masetto et al., 2012), Syagrus coronata (Mart.) Becc. (Porto et al., 2018), Euterpe precatoria Mart. (Costa et al., 2018) and Mauritia flexuosa L. f. (Almeida et al., 2018). In many species, the presence of light favors the germination of the seeds, while in others, the germinative behavior of the seeds is favored in the absence than in the presence of light (Melo et al., 2018). Thus, it can be inferred that both species of this research are classified as neutral photoblastics, since there was germination regardless of the presence or absence of light, being in accordance with what was proposed by Bewley et al. (2013) and Marcos Filho (2015).

It is observed that for the specie A. triandra the alternating temperature of 25-35 ºC reached higher values ​​for the characteristics germination percentage (76.76%) and GSI (0.2678), differing statistically from the others (Table 1). Germination in alternating and constant temperature regimes evidence the species' adaptation to the natural thermal fluctuations of the environment, which gives the seedlings greater ability to establish in the field, making them capable of withstanding adverse environmental conditions (Guedes et al., 2010). However, the difference between high and low temperatures must be equal to or greater than 10 ºC (Baskin & Baskin, 2014), thus simulating natural thermal variations. For Marcos Filho (2015) alternating temperatures favor, mainly, the germination of species that have not undergone an intense domestication process.

There are species that present higher percentages of germination when under alternating temperatures, and this is indicative of adaptive capacity to variations in the temperature of the environment (Martins et al., 2010), an event observed for the species A. vestiaria and A. triandra at temperatures of 20-30 and 25-35 ºC, respectively (Table 1). This is due to the similarity with the environmental germination where some species develop, in which the daytime temperatures are higher than the nighttime temperatures (Carvalho & Nakagawa, 2012). Additionally, knowledge about the response of seeds to temperature is essential, as it makes it possible to understand the range of tolerance of species in relation to temperature, as well as the climatic conditions in which crops can germinate and establish themselves properly (Motsa et al., 2015).

Table 1
Comparison of means for Germination percentage (G%) and germination speed index (GSI) of Areca vestiaria Giseke and Areca triandra Roxb seeds. ex Buch.-Ham. seeds, submitted to five temperatures and two light regimes

Averages	Areca vestiaria			Areca triandra
	G%1	GSI2		G%1	GSI2
DMSs (5%)	10.56	0.21		15.54	0.07
CV (%)	24.08	41.81		21.31	28.20
Light	30.74 a	0.3487 a		60.65 a	0.2067 a
Dark	29.75 a	0.3687 a		59.69 a	0.2043 a
DMSs (5%)	4.70	0.09		8.26	0.03
25 ºC	34.32 a	0.3703 ab		0.0 c	0.0 c
30 ºC	34.05 a	0.4397 a		53.62 b	0.1960 b
35 ºC	16.40 b	0.1805 b		56.83 b	0.1900 b
20-30 ºC	37.19 a	0.4583 a		53.49 b	0.1683 b
25-35 ºC	29.27 a	0.3447 ab		76.76 a	0.2678 a

• ¹
  Data transformed to arc sine (x/100)^1/2;
• ²
  Untransformed data; Means followed by the same letter do not differ in the column, by the Tukey Test, at 5% probability.
• DMSs (%): minimum significant difference and CV (%): coefficient of variation, expressed as a percentage.

Some species of palm trees also showed satisfactory results in the germination process of their seeds when under conditions of alternating temperatures, for example, Phoenix canariensis hort. ex Chabaud, 20-30 ºC (Pimenta et al., 2010); Copernicia alba Morong ex Morong & Britton, 20-30 ºC (Masetto et al., 2012); Bactris maraja Mart., 26-40 ºC (Rodrigues et al., 2014); Livistona rotundifolia (Lam.) Mart., 25-35 ºC (Viana et al., 2016) and Mauritia flexuosa L. f., 20-30 ºC (Almeida et al., 2018).

However, others germinated better at constant temperatures, such as Archontophoenix alexandrae H. Wendl. & Drude, 25 ºC (Teixeira et al., 2011); Oenocarpus bacaba Mart., 30 ºC (José et al., 2012); Sabal mauritiiformis (H. Karst.) Griseb. ex. H. Wendl., 30 ºC (Luz et al., 2014); Euterpe precatoria Mart., 20 ºC (Costa et al., 2018); Syagrus coronata (Mart.) Becc., 25 ºC (Porto et al., 2018).

The germination of the seeds of A. vestiaria (Figure 1) and A. triandra (Figure 2) responded heterogeneously to all temperatures in the conditions of presence and absence of light, with no expressive single peak, characterizing a polymodal behavior. The results may be related to the one proposed by Meerow and Broschat (2015), in which palm seeds in general have uneven and slow germination, caused by several factors, such as seed maturation stage, presence of mechanical dormancy caused by structures of the fruit, such as the rigid endocarp, which provide resistance to the embryo's expansion. The presence of the pericarp (epicarp and mesocarp) is a limiting factor in seed germination of some palm trees species (Teixeira et al.; 2011; Beckmann-Cavalcante et al. (2012); Pinto et al., 2012; Pereira et al., 2014). Based on these considerations, it was decided to remove these structures before starting the germination tests, in order not to interfere in the germination process.

Figure 1
Distribution of seed germination of Areca vestiaria Giseke submitted to five temperatures in the presence (A, B, C, D and E) and absence (F, G, H, I and J) of light.

Figure 2
Distribution of seed germination of Areca triandra Roxb. ex Buch.-Ham. submitted to four temperatures in the presence (A, B, C and D) and absence (E, F, G and H) of light.

For A. vestiaria, germination started on the 11^th day for all temperatures and light regimes (Figure 1). The stabilization in the presence of light occurred at 16^th; 43^rd; 44^th; 47^th and 52^nd day for temperatures of 35; 25; 30; 25-35 and 20-30 ºC, respectively. Larger peaks were noted on the 14^th; 15^th; 16^th; 24^th; 31^rst and 40^th for the temperature of 25 ºC with 3 germinated seeds (Figure 1A). For the temperature of 30 ºC, it occurred at 13^th and 28^th days with 5 seeds (Figure 1B). For 35 ºC it occurred on the 14^th day with 3 seeds (Figure 1C). The alternating temperature of 20-30 ºC reached higher peaks on the 13^th and 28^th days with 5 and 4 germinated seeds, respectively (Figure 1D), and 25-35 ºC on the 13^th day with 6 seeds (Figure 1E).

In the absence of light, germination stabilization for A. vestiaria occurred on th 28^th; 32^nd; 35^th and 48^th days for temperatures of 35; 20-30; 25-35; 25 and 30 ºC, respectively. Larger peaks were noted on the 22^nd day for a temperature of 25 ºC with 6 germinated seeds, 11^th (Figure 1F); 14^th; 15^th and 19^th day for 30 ºC with 4 seeds (Figure 1G). For 35 ºC, it occurred on the 11^th day with 5 seeds (Figure 1H), 20-30 ºC on the 19^th and 22^nd days with 6 seeds (Figure 1I), and 25-35 ºC on the 13^th and 17^th days, with 5 and 4 seeds (Figure 1J).

For the species A. triandra, in the presence of light, temperatures of 30 ºC and 20-30 ºC started the germination process on the 47^th day (Figure 2A; 2C), 35 ºC on the 49^th and 25-35 ºC on the 45^th day (Figure 2B; 2D). The germination stabilization occurred on the 96^th; 103^rd and 105^th day for temperatures of 25-35; 35; 30 and 20-30 ºC, respectively. Larger peaks were observed on days 63^rd; 65^th; 84^th and 96^th for temperatures of 35 ° C; 30 ºC; 20-30 ºC and 25-35 ºC with 9; 12; 6 and 16 germinated seeds, respectively.

In the absence of light, the germination process for A. triandra started on the 45^th day with an alternating temperature of 25-35 ºC (Figure 2H), after the 47^th day for 30; 35 and 20-30 ºC (Figure 2E; 2F; 2G). Stabilization occurred at 98^th days at 30 ºC and 25-35 ºC and at 105^th at 35 ºC and 20-30 ºC. With regard to the greatest germination peaks, these occurred at 63^rd; 96^th and 89^th days for temperatures of 30; 35 and 25-35 ºC, therefore the alternating temperature of 20-30 ºC reached peaks equal to 70^th, 84^th and 96^th days. The number of germinated seeds for each peak was 8; 10; 8 and 18, respectively, for 30; 35; 20-30 and 25-35 ºC. During the evaluations, the presence of fungi was observed, which possibly caused deterioration in the seeds, preventing and even delaying the germination process of the seeds that remained viable. Fungi cause decay and heating of the seed mass, resulting in an increase in the respiratory rate and production of mycotoxins, as pointed out by Carvalho & Nakagawa (2012). Thus, in the irrigations, distilled water with 0.2% nystatin was used to minimize contamination by fungi and were carried out whenever there was a need to replace water in the substrate, according to Luz et al. (2012).

The species Oenocarpus bacaba Mart. studied by José et al. (2012) required only 30 days for full germination. Viana et al. (2016) researching Livistona rotundifolia (Lam.) Mart., observed that this species needed 39 days. The species Euterpe oleracea Mart. it required 75 days, accordingly to Gonçalves et al. (2010), and Dypsis lutescens (H. Wendl.) Beentje & J. Dransf required 59 days (Pêgo & Grossi, 2016), corroborating with the exposed by Meerow and Broschat (2015).

The heterogeneity of germination within the same seed lot distributes the establishment of individuals over time and, with this, the population survives the phases inadequate to its development in the field (Carvalho & Nakagawa, 2012). However, the greater the number of days to emerge and the seedling to remain in the early stages of development, the greater the vulnerability to environmental conditions (Marcos Filho, 2015). However, for the producer who seeks to germinate quickly and homogeneously, it ends up becoming an obstacle, compromising his production, especially if it is on a large scale.

The results indicate that the species used in this research, even belonging to the same genus, have different mechanisms related to the germination process of their seeds, where A. vestiaria shows lower germination percentage compared to A. triandra. The temperature variation in the germination process of both species can be directly related to the breeding place of the matrices, as it is, according to Köppen (1948), for the region of Brumadinho, MG, average annual temperatures of 20.5 ºC and, for Jaboticabal, SP, of 32.5 ºC com. According to the same author, both regions have a humid subtropical climate, with a cold, dry winter and a hot, rainy summer.

For both species, varying temperatures (constant and alternating) to germination were shown, which may be indicative of adaptability to thermal fluctuations in the environment, thus making them capable of withstanding different climatic conditions. According to Silva et al. (2014), seeds of the same species can express diversified germinative behavior as a function of temperature and photoperiod, since within the same species there are variations between individuals as a function of the environment and their genetic constitution, corroborating the results obtained. However, the species A. triandra seems to present dormancy due to the prolonged period for the beginning of germination, thus requiring further studies dealing with the methods of overcoming dormancy, butdormancy but should always take into account its effective cost and ease of execution.

CONCLUSION

Temperatures of 25 ºC, 30 ºC, 25-35ºC and 20-30 ºC are indicated for germination of seeds of the species Areca vestiaria and, 25-35 ºC for Areca triandra, under the presence or the absence of light regimes, and are classified as neutral photoblastics.

Seed germination time varied for bothn species, with stabilization on the 52^nd day for Areca vestiaria and on the 105^th day for Areca trianda.

ACKNOWLEDGMENTS, FINANCIAL SUPPORT AND FULL DISCLOSURE

We would like to thank FCAV/UNESP Jaboticabal for supporting this work. This work was supported by the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) under a grant to the first author (number 148332/2019-6). There is no conflict of interest by the authors.

• 1
  Extracted from the first author’s Master Dissertation. Research financed with a scholarship from the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) to the first author.

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Publication Dates

• Publication in this collection
  17 Oct 2022
• Date of issue
  Sep-Oct 2022

History

• Received
  01 July 2021
• Accepted
  08 Dec 2021