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Article

Influence of Fruit Ripeness on Physiological Seed Quality of Maax Pepper (Capsicum annuum L. var. glabriusculum)

by
María Gabriela Dzib-Ek
1,
Rubén Humberto Andueza-Noh
2,*,
René Garruña
2,
Manuel Jesús Zavala-León
3,
Eduardo Villanueva-Couoh
1,
Benigno Rivera-Hernández
4,
Walther Jesús Torres-Cab
5,
Carlos Juan Alvarado-López
2 and
Roberto Rafael Ruíz-Santiago
6
1
División de Estudios de Posgrado e Investigación, Tecnológico Nacional de Mexico/Campus Conkal, Avenida Tecnológico s/n, Conkal C.P. 97345, Yucatán, Mexico
2
SECIHTI-Tecnológico Nacional de Mexico/Campus Conkal, Avenida Tecnológico s/n, Conkal C.P. 97345, Yucatán, Mexico
3
Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias, Campo Experimental Mocochá, Mocochá C.P. 97454, Yucatán, Mexico
4
Universidad Popular de la Chontalpa/División de Ciencias Básicas e Ingeniería, Carretera Cárdenas-Huimanguillo Km 2, Cárdenas C.P. 86529, Tabasco, Mexico
5
Instituto Tecnológico Superior de Hopelchén, Carretera Federal Campeche–Hopelchén, Km. 83, Hopelchén C.P. 24600, Campeche, Mexico
6
SECIHTI-Laboratorio Regional para el Estudio y Conservación de Germoplasma (GermoLab) del Centro de Investigación Científica de Yucatán, Parque Científico y Tecnológico de Yucatán, Km. 5.5, Carretera, Sierra Papacal-Chuburná Puerto, Sierra Papacal, Mérida C.P. 97302, Yucatán, Mexico
*
Author to whom correspondence should be addressed.
Agronomy 2025, 15(3), 747; https://doi.org/10.3390/agronomy15030747
Submission received: 20 February 2025 / Revised: 13 March 2025 / Accepted: 17 March 2025 / Published: 20 March 2025
(This article belongs to the Special Issue Seeds: Chips of Agriculture)
Browse Figures
Versions Notes

Abstract

:
Capsicum annuum L. var. glabriusculum is a semi-domesticated species of economic importance; however, its establishment in commercial plantations has been hampered by the low germination and emergence rates of its seeds. The aim of this study was to evaluate the effect of the fruit ripening stage on seed germination and seedling emergence in C. annuum var. glabriusculum. Seeds were extracted from fruits with six different ripening stages. The evaluated traits were the germination and emergence percentages, germination and emergence rates, and 17 physical traits of the seeds. According to the results, seeds extracted from red, orange, and pinto fruits presented better germination and seedling emergence percentages (85, 86, and 82% and 95, 93, and 94%, respectively). A principal component analysis showed that some differences in the physical traits of the seed were associated with the fruit ripening stages and seed development. A canonical discriminant analysis showed a high correlation between the fruit ripening stages and the physical and physiological characteristics of the seed, allowing the formation of four groups. The fruit ripening stages (pinto, orange, and red) influence the germination of the seeds and the emergence of the seedlings of C. annuum L. var. glabriusculum, so obtaining seeds from physiologically ripe fruits allows for obtaining seeds of better quality.

1. Introduction

The genus Capsicum includes 37 species, 5 of which have been domesticated and widely cultivated, including C. baccatum L., C. frutescens L., C. pubescens, C. chinense Jacq, and C. annuum L. Within each domesticated species there is a wide diversity that is reflected in the different sizes, shapes, colors, smells, flavors, and pungency levels of the fruits [1,2]. In Mexico, in addition to the five cultivated pepper species, we can find the wild species Capsicum annuum L. var. glabriusculum (Dunal) Heiser & Pickersgill, which has been identified as the progenitor and wild relative of the domesticated species C. annuum L. Capsicum annuum L. var. glabriusculum has a wide distribution throughout the Mexican Republic and in the south of the United States of America, which has allowed a great diversity of morphotypes to develop [3,4]. In Mexico, depending on the region where it grows, it is locally named pepper amashito, maax, piquín, chiltepín, or timpinchile, among other names. In this study, we will call it by the common name of the Mayan region, “chile maax” [3]. The maax pepper is important because it supports the economy of rural households who collect the fruit and sell it in local markets, it is part of Mexican gastronomy, and it is a plant genetic resource of great value [5]. The fruit in its immature stage is used for the artisanal production of sauces and pickles, while when ripe, it is dehydrated in the sun to be used as a condiment and to make sauces [6]. The fruit of the maax pepper is small, turns red when ripe, and is usually very spicy [7] according to the Scoville scale [8] C. annuum L. var. glabriusculum is considered the second hottest pepper in Mexico, as it varies between 100,000 and 200,000 SHU (Scoville units); it is considered a highly valuable plant genetic resource since it has been observed to present resistance to some viral groups, which makes it a candidate for use in genetic improvement programs as a source of adaptive traits for other species of cultivated peppers.
Currently, there is an interest in domesticating this species and establishing it as a commercial crop; for this reason, efforts have been made to promote its cultivation without much success, since one of the major limitations that this species presents is the low germination percentages of its seeds. This limitation is due to the fact that the seed presents physiological dormancy, according to which one or more internal conditions of the seed prevent it from germinating, even when it has optimal conditions for germination [9,10,11].
To break seed dormancy and improve the germination percentages of C. annuum L. var. glabriusculum, several studies have evaluated the application of pregermination treatments, but the results have not been very promising [12,13,14,15]. A strategy that has been successfully used to solve the problem of seeds with low germination percentages within the Capsicum genus has been the management of fruit ripening since it has been observed that harvesting fruits at different stages of ripening can influence the physical and physiological quality of the seed [16,17]. Hernández-Pinto et al. [18] reported that post-harvest storage of green and pinto fruits of Capsicum chinense Jacq. for 14 days increases the physiological quality of seeds and seedlings, presenting values similar to those of seeds obtained from mature fruits. Thus, the aim of this study was to evaluate the influence of the fruit ripening stage on the germination seed and seedling emergence of maax pepper (Capsicum annuum L. var. glabriusculum).

2. Materials and Methods

2.1. Experimental Site

The experiment was conducted on September 2023 at the Plant Physiology and Biotechnology Laboratory and experimental area of the Instituto Tecnológico de Conkal, located in Conkal, Yucatán, Mexico, at an altitude of 7 m and in an AW0 climate (warm subhumid with summer rains), with an average annual rainfall of 900 mm. The seeds were evaluated in a growth room with 22 ± 1 °C and 30% relative humidity and in field conditions where the average daily temperature was 32 ± 2 °C with 60% relative humidity.

2.2. Plant Material

The seeds of maax pepper (Capsicum annuum L. var. glabriusculum) were obtained from the Universidad Popular de la Chontalpa. The seeds were extracted from fruits of six different ripeness stages, as follows: (1) green—immature fruits, harvested 25 days post-anthesis; (2) olive—olive-green fruits, harvested 35 days post-anthesis; (3) pinto—olive-green to orange fruits, harvested 42 days post-anthesis; (4) orange—ripe fruits, harvested 49 days post-anthesis; (5) red—fully ripe fruits harvested 56 days post-anthesis; and (6) overripe—dehydrated fruits, harvested 70 days post-anthesis (Figure 1).
The viable seeds from each lot were selected using the immersion method, which consisted of immersing the seeds in water for 2 min. The seeds that floated were considered non-viable or empty seeds, and seeds that did not float were considered viable. Seeds were disinfected with a 2% sodium hypochlorite solution for 3 min, rinsed with running water under the tap, and then given a final rinse with distilled water. After, the seeds were dried in mesh bags at room temperature (30 °C ± 2) for 72 h. Once dried, they were stored in airtight plastic bags and kept at 10 °C for 6 months until their evaluation.

2.3. Variables Evaluated

The physiological and physical variables of the seeds were evaluated. The physiological variables included the moisture content, electrical conductivity, seed germination rate and percentage, and seedling emergence rate and percentage.
The seed moisture content (%H) was measured using a forced-air convection furnace (TERLAB TE-H80DM) at 105 °C for 24 h. Three replicates of 0.120 g (100 seeds) were used for each seed lot.
The seed’s electrical conductivity (EC) was measured using a conductivity meter (Consort C931, Debruyne Instruments, Wichelen, Belgium). Three replicates of 50 seeds per lot were used; each replicate was immersed in 50 mL of deionized water with electrical conductivity and incubated for 24 h at 25 °C [19]. After incubation, the electrical conductivity of the solution was measured and recorded.

2.4. Germination Test and Germination Rate Index

Germination tests were carried out for each seed lot, with four replications of 25 seeds, resulting in a total of 100 seeds evaluated per lot for each of the six fruit maturation stages. The previously disinfected seeds were placed in 100 mm in diameter × 15 mm in height Petri dishes containing two layers of absorbent paper moistened to 2.5 times their dry weight. The Petri dishes were then placed in a growth chamber under a controlled temperature (25 ± 1 °C), constant humidity (70%), and total darkness. The number of germinated seeds was recorded every 3rd day over a 28-day period. The seeds were considered germinated when root protrusion was observed. From the data collected, the germination percentage (% G) and germination rate index (GSI) were calculated as follows:
Germination percentage (% G) = [(Number of seeds germinated)/(Number of seeds sown)] × 100
Germination rate index (GSI) = ∑[ni/ti]
where ni is the number of seeds germinated in the time interval ti, and ti is the period in days from sowing to the final day of the experiment [20].

2.5. Emergency Test and Emergency Rate Index

For the seed emergence test, polystyrene trays with 200 cavities were used, previously disinfected with 6% sodium hypochlorite, and filled with Cosmopeat® substrate (fine-fiber peat moss, vermiculite, wetting agents, and nutrients). One seed was placed in each cavity at a depth of approximately 0.5 cm. Four replicates of 25 seeds from each seed lot were sown, with a total of 100 seeds per lot. The trays were then covered with black plastic and maintained at a temperature of 25 °C. The trays were kept moist by watering as needed to meet the seedlings’ requirements. Emergence was monitored daily, and a seedling was considered emerged when its cotyledons were perpendicular to the upright hypocotyl. Using the data obtained after 28 days, the seedling emergence percentage (%E) and seedling emergence rate index (ESI) were calculated according to Maguire [21].
Percentage of seedling emergence (% E) = [(No. of seedlings emerged)/(No. of seeds sown)] × 100
Seedling emergence rate index (ESI) = ∑[ni/ti]
where ni is the number of seedlings that emerged in the time interval ti, and ti is the period in days from sowing to the final day of the experiment [20].

2.6. Evaluation of the Physical Traits of the Seed

To evaluate the physical traits of the seed, four replicates per seed lot were used, each replicate containing 25 seeds, for a total of 100 seeds for each lot. The seeds were arranged on the scanner bed, and the area occupied by the seeds was covered with a tray featuring a blue background to capture digital images. The images were saved as JPG files, and the physical traits of the seeds were then measured using a tool for seed image analysis [22]. The physical seed traits were the area, perimeter, convex area, convex perimeter, feret, breadth, compactness, solidity, thinness, concavity, roundness, sphericity, irregularity, endocarp, circularity, rectangularity ratio (AspRatio), and average length of the ratio (AvgR). All measurements were taken using a tool for seed image analysis [22].

2.7. Experimental Design and Statistical Analysis

The experimental design applied in the germination tests and the seedling emergence test, and in the evaluation of the physical traits of the seeds, was completely randomized. Each seed lot was treated as an independent treatment, with four replicates per treatment in each test. The data for each variable were tested for normality using the Shapiro–Wilk test and were subjected to an analysis of variance (ANOVA) at a 95% confidence level. Where significant differences were found, means were compared using the Tukey test (p ≤ 0.05) with INFOSTAT software version 2020e.
The physical characteristics were analyzed using multivariate statistics through principal component analysis (PCA). A preliminary analysis, which included all the physical variables, showed that 13 variables contributed the most to the variance (area, perimeter, convex area, convex perimeter, feret, breadth, compactness, thinnessR, roundness, sphericity, circularity, rectangularity ratio (AspRatio), and average length of the ratio (AvgR)). Thus, the final PCA was based on these 13 variables. To examine the relationship between the physiological and physical seed traits and the fruit ripening stages, a canonical discriminant analysis (CDA) was performed. Both the PCA and CDA analyses were conducted using R statistical software version 4.3.2.

3. Results

3.1. Physiological Parameters of Maax Pepper Seeds (Capsicum annuum L. var. glabriusculum)

The moisture content (%) of the seed lots at different fruit ripening stages is shown in Figure 2a. No significant differences (p ≤ 0.05) were observed in moisture content across the different treatments (green, olive, pinto, orange, red, overripe). The moisture content ranged from 6 to 11%, with an average of 8.6%.
The electrical conductivity (EC) of the seed lots at different fruit ripening stages showed significant differences (p ≤ 0.05) among the treatments (Figure 2b). The mean comparison revealed that seeds at the pinto ripening stage exhibited the lowest electrical conductivity, which was statistically similar to that in the seeds obtained from fruit at the ripening stages of orange, red, and overripe. The seeds of fruit in the olive ripening stage had the highest electrical conductivity, similar to the seeds from fruits in the green ripening stage (Figura 2b).
The germination percentages (%G) of the seed lots at different fruit ripening stages showed significant differences (p ≤ 0.05) among the treatments (Figure 2c). Seed lots extracted from fruits in the olive, pinto, orange, and red ripening stages showed the highest germination, presenting values of 75, 82, 86, and 85%, respectively. The seeds from the green stage did not germinate during the germination period, and seeds from the overripe ripening stage showed 57% germination (Figure 2c).
The percentage of seedling emergence was affected by the stage of fruit ripening, the seed lots from the green and overripe stages being the ones that obtained the lowest values of seedling emergence, at 22 and 61%, respectively. Seeds extracted from olive, pinto, orange, and red fruit ripening stages showed the best seedling emergence response, presenting values of 92, 94, 93, and 95%, respectively (Figure 2d).
The seeds extracted from fruits in the red, orange, and pinto ripening stages showed significant statistical differences in seed germination rate, with five seeds germinating per day, while seeds extracted from the green stage did not germinate. The seedling emergence rate was higher in seed lots extracted from fruits with olive, pinto, and orange ripening stages, with ten, nine, and eight emerged seedlings, respectively, and the lowest results were presented by seed lots obtained from fruits in the green and overripe stages (Table 1).

3.2. Morphological Traits of Maax Pepper Seeds (Capsicum annuum L. var. glabriusculum)

A principal component analysis (PCA) revealed that the first two principal components (PC) explained 82.4% of the total accumulated variance. PC1 contributed 55.8% to the total variation, while PC2 explained 26.6% of the total accumulated variation. The scatter plot based on these two principal components showed differences in the physical characteristics of each seed lot, separating the seed lots (Figure 3). The seed lot extracted from fruits at the orange ripening stage was characterized by its area, perimeter, convex perimeter, breath, average length of the ratio (AvgR), and seed feret. The seed lot obtained from the overripe stage featured roundness, compactness, and sphericity in its seeds. The olive stage was distinguished for its seeds’ thinnessR and convexArea. Seeds from the pinto stage were characterized by circularity in their shape, while the red-stage seeds had a higher rectangularity ratio (AspRatio). Finally, the seed lot from the green stage was characterized by the smallest average radius, reflecting the smallest seed size.

3.3. Relationship Between Physical and Physiological Traits of Seed and Ripening Stages of Maax Pepper Fruit (Capsicum annuum L. var. glabriusculum)

A canonical discriminant analysis (CDA) revealed that the correlations between the physical and physiological seed traits and fruit ripening stages of the six seed lots were highest for the first two canonical axes (0.99 and 0.99, respectively; Table 2). Together, both these axes explained 96.5% of the total accumulated variation (82.4% and 14.1% for variables Can 1 and Can 2, respectively; Table 2). Figure 4a,b show that in the first canonical axis (Can 1), morphological traits such as seed area (AR), seed perimeter (PR), seed feret (LR), and seed diameter (AN) contributed the most to the observed variation. These traits were primarily associated with the seed lots extracted from the olive, overripe, and orange fruit ripening stages. In the second canonical axis (Can 2), the physiological variables germination (GR), seedling emergence (EM), germination rate (GR), and emergence rate (ER) contributed most significantly to the observed variation. These traits were more closely related to the seed lots obtained from fruits with red, pinto, orange, and olive ripening stages (Figure 4c,d).
The scatter diagram of the first two canonical axes (Figure 5) illustrates that the six evaluated seed lots form four groups based on their physical and physiological traits. Group I consists of seed lots derived from fruits at the pinto and red ripening stages. This group is characterized by a higher germination percentage, higher germination rate, and larger seed convex area. Group II includes seed lots extracted from fruits at the olive and orange ripening stages. These seed lots exhibit a higher seedling emergence percentage, higher seedling emergence rate, and thinner seeds. Group III comprises the seed lot obtained from fruits at the overripe stage, distinguished by superior performance in the physical traits of the seed. Group IV comprises the seed lot extracted from fruits in the green ripening stage and is characterized by the lowest values in the physiological variables, as well as being related to the physical traits of circularity and convex perimeter.

4. Discussion

4.1. Moisture Content and Electrical Conductivity of Maax Pepper (Capsicum annuum L. var. glabriusculum)

The results for the seed moisture content show a humidity range between 6 and 11%, which indicates that the seeds are alive. According to SNICS [23], the moisture content of the seed observed in the six evaluated seed lots is within the acceptable range established in all seed categories (basic, registered, certified, and enabled). On the other hand, taking into account that no significant differences were observed in the seed moisture of all the evaluated seed lots, we can conclude that the seed lots from the different stages of fruit ripening were under the same moisture conditions at the beginning of the germination and emergence tests, which allowed us to obtain consistent results.
The electrical conductivity (EC) test is used as an indicator of seed viability [20]. By measuring the EC of the imbibition solution, the amount of solutes released by the seeds (amino acids and electrolytes) could be determined; this amount is inversely proportional to the seeds’ viability. A higher EC indicates that the organization or integrity of the seed’s cell membranes is deficient and that the seed will therefore present viability problems; on the other hand, low levels of electrical conductivity indicate greater integrity of the seeds’ cell membrane, further indicating better seed viability [18]. In this way, Ayala et al. [24] reported that a decrease in the EC increased the germination of Capsicum seeds. However, in this study, no relationship was observed between germination and EC. Based on the results, it is observed that EC is not the main factor influencing germination in this variety. Since the EC of the green stage was statistically similar to the more advanced stages of maturity, however, the green stage did not germinate. The electrolyte that is released and increases the EC is likely not directly related to germination. However, studies would have to be carried out specifically on this variety.

4.2. Seed Germination and Seedling Emergence of Maax Pepper (Capsicum annuum L. var. glabriusculum)

In this study, the seed lot extracted from fruit in the green ripening stage negatively influenced the number of germinated seeds when compared to the seed lots extracted from fruits in the most advanced of ripening stages (olive, pinto, orange, and red ripening stages), where the germination percentages were the highest. This result can be attributed to the fact that the seeds obtained from fruits in the green ripening stage did not reach complete physiological maturity and therefore had immature or poorly developed embryos, while the better germination performances of seeds obtained from fruits in an advanced stage of maturity can be attributed to the fact that the seeds reached physiological maturity and therefore possessed physiologically mature embryos that provided an advantage in germination. These results are similar to those reported by Hernández et al. [18], who evaluated the effects of different fruit ripening stages on the physiological quality of habanero pepper seeds (Capsicum chinense Jacq). On the other hand, our results are superior to those reported by Brondo-Ricárdez et al. [15], who evaluated the effects of pregermination treatments on the physiological quality of maax pepper seeds (Capsicum annuum L. var. glabriusculum) obtained from mature fruits. Similar results have also been observed in different cucurbit species, where the best germination percentages were observed in seeds extracted from fully ripe fruits [25,26].
Regarding the seed lot extracted from fruits in the overripe stage, the germination percentage was low. This could be due to the fact that harvesting fruits too late can increase the risk of seed deterioration and thereby influence the quality of the seeds [27]. This result agrees with those of Sripathy and Groot [28], who pointed out that during the stages of a seed’s development, its physiological quality evolves until it reaches maximum physiological maturity; subsequently, its deterioration begins due to the effects of aging. Regarding the results obtained for the emergence of seedlings, a pattern similar to that observed in seed germination was registered, due to the fact that the seeds extracted from fruits with more advanced ripening stages (seeds extracted from fruits in the olive, pinto, orange, and red ripening stages) showed the highest seedling emergence values, and seeds extracted from fruits in the green maturity stage showed the lowest seedling emergence percentages, which means that the seeds had not fully matured.
Similar results were presented by Santamaria and Zavala [17], who reported that extracting seeds from fruits in more advanced stages of ripening (overripe) results in seeds of lower quality. These same authors observed, when evaluating different stages of fruit ripening and the effects of their relationship with the storage period on the germination of Capsicum annuum L. seeds, that the lowest percentage of emerged seedlings was displayed by seeds extracted from physiologically immature fruits, indicating that these fruits do not produce seeds of good physiological quality. In contrast, seeds obtained from completely mature fruits have the capacity to produce seeds of better physiological quality, thereby allowing the development of better-quality seedlings. These authors emphasized that seeds from mature fruits have reached adequate physiological maturity, providing the necessary attributes for superior performance in the field and ensuring higher productivity. Similarly, Valdez-Eleuterio [29] highlighted that evaluating seed germination, seedling emergence, and vigor in an environment similar to the crop’s natural one offers more realistic insights into the quality of seed lots. This approach also enables the assessment of their true potential under field conditions. Criollo and Upegui [30], in a study on the physiological quality of Physalis peruviana seeds, reported findings consistent with those of this study; they observed the lowest seedling emergence values in seeds obtained from immature fruits and the best emergence percentages in seeds extracted from mature fruits. The authors emphasized that seeds from mature fruits attained adequate physiological maturity, and they also showed the necessary conditions to achieve greater performance potential in the field, ensuring greater productivity.

4.3. Seed Germination Rate and Seedling Emergence Rate of Maax Pepper (Capsicum annuum L. var. glabriusculum)

The seed germination rate was higher in seed lots extracted from fruits in the pinto, orange, and red ripening stages. This result agrees with those of Sanches et al. [31], who evaluated the effects of fruit maturity on the physiological quality of the seed of Diospyros inconstans Jacq, observing that the seeds of red fruits showed a higher germination rate. This result may be associated with the maturation of the seeds since red fruits are in their physiological stage of maturity. In the same area, Silva et al. [32] obtained similar results in seeds of Trichosanthes cucumerina L. (Cucurbitaceae), reporting that the germination rate became higher with further delay in the fruit harvest time. Ayala-Villegas et al. [24] mentioned that the best physiological quality was observed in the seed germination kinetics of Capsicum annuum L. It was reached when the seeds were extracted 15 days after the fruit was harvested, when the fruits were completely red, while in the Guajillo pepper, the maximum speed of germination was achieved when the fruits were harvested at the beginning of the fruit color change and the seeds were extracted 15 days after the harvest; therefore, these authors concluded that harvesting fully ripe fruits or fruits at the beginning of physiological maturity and extracting the seed 15 days after harvest improves the physiological quality of Capsicum annuum seed.
The seedling emergence rate defines the plant’s performance capacity in the field, and this performance capacity is improved by the use of good-quality seeds; that is, vigorous seeds will give rise to vigorous and more productive plants [22]. This effect could be observed in the results of our study since the seeds from the most advanced stages of fruit maturity achieved a higher emergence rate, which indicates that the seed was able to reach physiological maturity and therefore showed greater vigor. Hernández-Pinto et al. [18] reported that the highest seedling emergence rate for Capsicum chinense Jacq variety Mayapán was obtained from seeds extracted from mature fruits stored for 14 days. The results regarding the seedling emergence rate in this study coincide with the results of studies carried out by Neto et al. [25], who reported that, in melon (Cucumis melo L.), the emergence rate was higher in seeds extracted from mature fruits, compared to seeds extracted from green fruits. Depending on the stage of maturation and the emergence of seedlings, these parameters are likely to be related to physiological maturity and the accumulation of reserves in the seeds. Santos et al. [33] evaluated the seedling emergence rates of seed lots extracted at different stages of maturation of pitahaya (Hylocereus spp.) and obtained percentage emergence values ranging from 34% to 96%. An important factor to take into account in relation to the seedling emergence rate is the type of substrate used in the test, since those substrates that provide adequate porosity—that is, that facilitate the supply of oxygen and water retention—help in improving the speed of seedling emergence.

4.4. Physical Traits of Maax Pepper Seeds (Capsicum annuum L. var. glabriusculum)

In this study, the seed lots obtained from fruits in the pinto, orange, and red ripening stages exhibited better morphological traits compared to the seed lots obtained from fruits in the green ripening stage, which were smaller in size. This suggests that the seeds obtained from green fruits did not fully complete their physiological maturation and exhibited less seed filling compared to those from more mature fruits (fruits in pinto, orange, and red ripening stages). This finding is consistent with those of Cano-Vázquez et al. [4], who observed that the physical traits of the piquín pepper seed (Capsicum annuum L. var. glabriusculum), such as area, perimeter, diameter, and roundness, were directly related to embryo size, and concluded that larger seeds had larger embryos, resulting in better physiological attributes that support more successful germination and emergence. Similarly, Martínez et al. [34] noted that seeds with greater area, length, and weight in Capsicum annuum L. exhibited higher germination and emergence percentages. In contrast, Quevedo and Laurentin [35] found no variation in seed physical attributes such as the color, surface, size, and number of seeds per fruit in sweet pepper (Capsicum chinense Jacq.). In the case of Jatropha curcas L., it was observed that seeds obtained from green fruits had a lower tissue density and presented a greater number of malformed seeds compared to seeds extracted at more advanced stages of maturity [36].

4.5. Relationship Between the Stage of Fruit Ripening, Physical, and Physiological Traits of the Seed of Maax Pepper (Capsicum annuum L. var. glabriusculum)

The results of the canonical discriminant analysis showed that the seeds extracted from fruits at more advanced ripening stages (pinto and red) presented better physiological seed quality. This can be attributed to the concurrent maturation of seeds with fruit ripening. At advanced fruit ripening stages, seeds complete embryo development and achieve maximum dry matter accumulation, marking the conclusion of the seed-filling phase. Consequently, the fruit’s maturity stage at harvest is a critical factor influencing seed quality [28].
On the other hand, extracting seeds from fruits at early ripening stages (green fruits) can result in poor physiological seed quality due to the incomplete development of essential structures and protective mechanisms of the embryo and seed [37]. The results of this research coincide with those reported by Martínez et al. [34], who observed that seeds extracted from physiologically mature fruits of huacle chili (Capsicum annuum L.) demonstrated superior physical and physiological quality. Similarly, Dos Santos et al. [27] determined that the maturity of Capsicum chinense Jacq. fruits prior to seed extraction significantly affect embryo development. Thus, extracting seeds from mature fruits results in higher germination rates and seed vigor. Pinheiro et al. [36] reported that there is a relationship between the physiological potential of J. curcas seeds and the different stages of maturity and physical attributes, indicating that seeds extracted from yellow and brownish-yellow fruits have a higher physiological quality. Similar results were derived by Medeiros et al. [38], who reported that, in seeds of Leucaena leucocephala, physical traits related to seed shapes, such as perimeter and circularity, are efficient indicators of seedling performance, such that the larger the perimeter and the lower the circularity, the better the seedling performance observed.

5. Conclusions

The seeds of Capsicum annuum L. var. glabriusculum obtained from fruits in the pinto, orange, and red ripening stages presented better physical and physiological quality. The seeds from the green ripening stage showed higher electrical conductivity, lower physical characteristics, and lower physiological quality. Seeds extracted from harvested fully ripe fruits of Capsicum annuum L. var. glabriusculum showed better germination performance and seedling emergence. Therefore, obtaining seeds from physiologically mature fruits allows for a higher yield and better seed quality. In Capsicum annuum L. var. glabriusculum, the ripening stage of the fruit should be considered an important criterion for deriving good-quality seeds.

Author Contributions

Conceptualization, R.H.A.-N., M.G.D.-E. and R.G.; methodology, M.G.D.-E., W.J.T.-C., R.R.R.-S. and E.V.-C.; software, R.H.A.-N., M.G.D.-E. and R.G.; validation, R.H.A.-N., M.J.Z.-L., E.V.-C., C.J.A.-L. and B.R.-H.; formal analysis, M.G.D.-E., M.J.Z.-L., W.J.T.-C. and R.R.R.-S.; investigation, R.H.A.-N., M.G.D.-E. and B.R.-H.; resources, E.V.-C., C.J.A.-L. and B.R.-H.; writing—original draft preparation, R.H.A.-N. and M.G.D.-E.; writing—review and editing, R.H.A.-N., R.G. and M.J.Z.-L.; visualization, M.G.D.-E., W.J.T.-C. and C.J.A.-L.; supervision, R.H.A.-N., R.G., E.V.-C. and B.R.-H. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Data Availability Statement

The data presented in this study are available on request from the corresponding author.

Acknowledgments

The first author thanks the Secretariat of Science, Humanities, Technology and Innovation from Mexico (SECIHTI) for their graduate student scholarship number: 703490.

Conflicts of Interest

The authors declare no conflicts of interest.

References

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Agronomy 15 00747 g001
Figure 1. Ripening stages of maax pepper fruits (C. annuum L. var. glabriusculum) from which the six lots of seeds were extracted: 1 green (25 days post-anthesis (dpa)), 2 olive (35 dpa), 3 pinto (42 dpa), 4 orange (49 dpa), 5 red (56 dpa), and 6 overripe (70 dpa).
Figure 1. Ripening stages of maax pepper fruits (C. annuum L. var. glabriusculum) from which the six lots of seeds were extracted: 1 green (25 days post-anthesis (dpa)), 2 olive (35 dpa), 3 pinto (42 dpa), 4 orange (49 dpa), 5 red (56 dpa), and 6 overripe (70 dpa).
Agronomy 15 00747 g001
Agronomy 15 00747 g002
Figure 2. Physiological variables of seeds extracted from fruits with different ripening stages of maax pepper (Capsicum annuum L. var. glabriusculum): (a) moisture content, (b) electrical conductivity, (c) germination, and (d) seedling emergence. Different letters indicate statistically significant differences between treatments (Tukey, p ≤ 0.05).
Figure 2. Physiological variables of seeds extracted from fruits with different ripening stages of maax pepper (Capsicum annuum L. var. glabriusculum): (a) moisture content, (b) electrical conductivity, (c) germination, and (d) seedling emergence. Different letters indicate statistically significant differences between treatments (Tukey, p ≤ 0.05).
Agronomy 15 00747 g002
Agronomy 15 00747 g003
Figure 3. Scatter plot of seeds extracted from fruits with different ripening stages of maax pepper (Capsicum annuum L. var. glabriusculum) based on the first two principal components using 13 seed morphological traits. Different letters indicate statistically significant differences between treatments (Tukey, p ≤ 0.05).
Figure 3. Scatter plot of seeds extracted from fruits with different ripening stages of maax pepper (Capsicum annuum L. var. glabriusculum) based on the first two principal components using 13 seed morphological traits. Different letters indicate statistically significant differences between treatments (Tukey, p ≤ 0.05).
Agronomy 15 00747 g003
Agronomy 15 00747 g004
Figure 4. Relationship between physical and physiological variables of seeds extracted from fruits with different ripening stages of Capsicum annuum L. var. glabriusculum. (a,b) First canonical function (Can 1). (c,d) Second canonical function (Can 2). AR (area), PR (perimeter), LR (feret), AN (breadth), DZ (thinness), RZ (roundness), AC (convex area), PR (convex perimeter), CR (circularity), GR (germination), EM (emergence), TG (germination rate), TE (emergence rate), and CE (electrical conductivity).
Figure 4. Relationship between physical and physiological variables of seeds extracted from fruits with different ripening stages of Capsicum annuum L. var. glabriusculum. (a,b) First canonical function (Can 1). (c,d) Second canonical function (Can 2). AR (area), PR (perimeter), LR (feret), AN (breadth), DZ (thinness), RZ (roundness), AC (convex area), PR (convex perimeter), CR (circularity), GR (germination), EM (emergence), TG (germination rate), TE (emergence rate), and CE (electrical conductivity).
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Figure 5. Scatter diagram of canonical discriminant analysis showing the influence of physical variables of seeds extracted from fruits of maax pepper (Capsicum annuum L. var. glabriusculum) in different ripening stages. AR (area), PR (perimeter), LR (feret), AN (breadth), DZ (thinness), RZ (roundness), AC (convex area), PC (convex perimeter), CR (circularity), GR (germination), EM (emergence), TG (germination rate), TE (emergence rate), CE (electrical conductivity).
Figure 5. Scatter diagram of canonical discriminant analysis showing the influence of physical variables of seeds extracted from fruits of maax pepper (Capsicum annuum L. var. glabriusculum) in different ripening stages. AR (area), PR (perimeter), LR (feret), AN (breadth), DZ (thinness), RZ (roundness), AC (convex area), PC (convex perimeter), CR (circularity), GR (germination), EM (emergence), TG (germination rate), TE (emergence rate), CE (electrical conductivity).
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Table 1. Seed germination and seedling emergence rates of six seed lots of maax pepper (Capsicum annuum L. var. glabriusculum) extracted from fruit in different ripening stages. Different letters indicate statistically significant differences among treatments (Tukey, p ≤ 0.05).
Table 1. Seed germination and seedling emergence rates of six seed lots of maax pepper (Capsicum annuum L. var. glabriusculum) extracted from fruit in different ripening stages. Different letters indicate statistically significant differences among treatments (Tukey, p ≤ 0.05).
TreatmentsGermination Rate
(Seeds Germinated/Day)		Emergency Rate
(Seedlings Emerged/Day)
Green0 ± 0.00 c0.28 ± 0.01 d
Olive4.68 ± 0.44 ab10.87 ± 0.65 a
Pinto5.07 ± 0.09 a8.08 ± 0.80 ab
Orange5.28 ± 0.26 a9.36 ± 0.093 ab
Red5.67 ± 0.22 a6.45 ± 0.66 bc
Overripe3.40 ± 0.22 b2.91 ± 0.40 cd
Table 2. Summary of the canonical discriminant analysis of the physical and physiological traits of seeds of Capsicum annuum L. var. glabriusculum. **, *** = highly significant differences (p ≤ 0.01; 0.001).
Table 2. Summary of the canonical discriminant analysis of the physical and physiological traits of seeds of Capsicum annuum L. var. glabriusculum. **, *** = highly significant differences (p ≤ 0.01; 0.001).
Canonical VariableCanonical CorrelationSelf-ValueEigenvalue RatioCumulative ProportionProbability Value
Can 10.998801.29682.38982.3896 × 10−11 ***
Can 20.992137.23414.11096.5002 × 10−06 ***
Can 30.96730.1093.09599.5960.005183 **
Can 40.7813.5740.36799.9630.441521
Can 50.2620.3560.036100.0000.954468
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Dzib-Ek, M.G.; Andueza-Noh, R.H.; Garruña, R.; Zavala-León, M.J.; Villanueva-Couoh, E.; Rivera-Hernández, B.; Torres-Cab, W.J.; Alvarado-López, C.J.; Ruíz-Santiago, R.R. Influence of Fruit Ripeness on Physiological Seed Quality of Maax Pepper (Capsicum annuum L. var. glabriusculum). Agronomy 2025, 15, 747. https://doi.org/10.3390/agronomy15030747

AMA Style

Dzib-Ek MG, Andueza-Noh RH, Garruña R, Zavala-León MJ, Villanueva-Couoh E, Rivera-Hernández B, Torres-Cab WJ, Alvarado-López CJ, Ruíz-Santiago RR. Influence of Fruit Ripeness on Physiological Seed Quality of Maax Pepper (Capsicum annuum L. var. glabriusculum). Agronomy. 2025; 15(3):747. https://doi.org/10.3390/agronomy15030747

Chicago/Turabian Style

Dzib-Ek, María Gabriela, Rubén Humberto Andueza-Noh, René Garruña, Manuel Jesús Zavala-León, Eduardo Villanueva-Couoh, Benigno Rivera-Hernández, Walther Jesús Torres-Cab, Carlos Juan Alvarado-López, and Roberto Rafael Ruíz-Santiago. 2025. "Influence of Fruit Ripeness on Physiological Seed Quality of Maax Pepper (Capsicum annuum L. var. glabriusculum)" Agronomy 15, no. 3: 747. https://doi.org/10.3390/agronomy15030747

APA Style

Dzib-Ek, M. G., Andueza-Noh, R. H., Garruña, R., Zavala-León, M. J., Villanueva-Couoh, E., Rivera-Hernández, B., Torres-Cab, W. J., Alvarado-López, C. J., & Ruíz-Santiago, R. R. (2025). Influence of Fruit Ripeness on Physiological Seed Quality of Maax Pepper (Capsicum annuum L. var. glabriusculum). Agronomy, 15(3), 747. https://doi.org/10.3390/agronomy15030747

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