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Vol. 9(6), pp. 627-635, 6 February, 2014
DOI:10.5897/AJAR2013.7133
ISSN 1991-637X ©2014 Academic Journals
http://www.academicjournals.org/AJAR
African Journal of Agricultural
Research
Full Length Research Paper
Physiological and biochemical alterations during
germination and storage of habanero pepper seeds
Franciele Caixeta*, Édila Vilela De Resende Von Pinho, Renato Mendes Guimarães,
Pedro Henrique Andrade Rezende Pereira and Hugo Cesar Rodrigues Moreira Catão
Department of Agriculture, Federal University of Lavras, C.P. 3037, 37200-000, Lavras, MG, Brazil.
Accepted 12 November, 2013
The objective of this study was to evaluate physiological and biochemical alterations during the
development and storage of habanero pepper seeds with a view toward determining the time of harvest.
Seeds were manually extracted from the fruit at three stages of development: E1 (fruit with first signs of
yellowing), E2 (mature fruit) and E3 (mature fruit submitted to seven days of rest). After drying, seeds
with 8% water content were stored at 10°C for 0, 4 and 8 months, and their quality evaluated by means
of germination and vigor tests. Activities of the enzymes α-amylase, endo-β-mannanase, esterase,
Superoxide Dismutase (SOD), malate dehydrogenase (MDH) and alcohol dehydrogenase (ADH) were
evaluated during germination at 0, 48, 96 and 144 h after seeding. A randomized block design was used
in a 3 × 3 factorial design (stages of development × storage) with 4 replications. Lower germination and
vigor values were observed for the E1 stage seeds at all storage periods. In recently stored seeds,
greater germination and vigor values were observed for the E3 stage seeds. Dormancy was observed
principally in recently stored seeds and this was overcome at four months of storage. In summary, the
physiological tests and activity of the enzymes evaluated indicated that the habanero pepper should be
harvested at the E3 stage for a higher seed quality.
Key words: Capsicum chinense, seed maturation, seed quality.
INTRODUCTION
The demand for high quality seeds has grown
substantially in recent years, which requires that seed
companies adopt advanced technologies during
production, processing and storage processes. The
adoption of new seed technologies requires knowledge of
the factors that influence physiological quality on the one
hand and on the physiological and biochemical
alterations that occur during the germination and storage
processes on the other hand. Studies related to
maturation and harvest of seeds is important since seeds
reach their maximum quality in the field. Such knowledge
is necessary for seed producers to determine the ideal
time for harvest that would minimize seed quality
deterioration caused by a prolonged period in the field,
and increase seed production, by preventing a too early
harvest since this may result in a large proportion of
immature seeds (Vidigal et al., 2009). In species with
fleshy fruit, it has been observed that seeds maintained
for a certain period of time in the fruit after harvest
complete the maturation process, reaching maximum
levels of germination and vigor (Barbedo et al., 1994;
Vidigal et al., 2006; Dias et al., 2006). Therefore, post-
harvest storage of the fruit before seed extraction may be
advantageous because it allows early harvest, and avoid
exposure to possible unfavorable conditions that may
deteriorate seed quality (Barbedo et al., 1994;
*Corresponding author. E-mail: francielecaixeta@yahoo.com.br
628 Afr. J. Agric. Res.
Table 1. Mean values of maximum (Tx), minimum (Tn), and mean (Tm) temperatures, relative humidity (UR), total rainfall (Pt) and insolation
(I) of the months corresponding to the crop development period of the 2007 harvest.
Months
Tx (ºC)
Tn (ºC)
Tm (ºC)
UR (%)
Pt (mm)
I (h)
January
27.6
18.7
21.9
85.7
554.7
3.1
February
28.9
18.1
22.6
73.1
151.3
7.5
March
30.9
18.1
23.6
66.5
35.4
9.0
April
28.1
17.2
21.8
72.4
35.6
7.3
May
25.7
12.8
18.1
70.6
30.4
7.5
June
25.8
11.1
17.3
66.3
5.9
8.8
July
25.4
11.1
17.1
66.8
17.6
7.7
Source: Agrometeorology Sector of the Engineering Department – UFLA.
Dias et al., 2006).
Deterioration is a process determined by a series of
physiological, biochemical, physical and cytological
alterations that occur in a progressive manner, leading to
lower quality and culminating in death of the seed
(Freitas, 2009). The main alterations related to the
deterioration process are degradation and inactivation of
enzymes, reduction of respiratory activity and loss of
integrity of cellular membranes (Copeland and McDonald,
2001). The speed of this deterioration process is
influenced by storage conditions. During seed storage, it
is necessary to maintain adequate temperature and
humidity conditions in the attempt to preserve quality. In
this research, physiological and biochemical alterations
during development of yellow habanero pepper
(Capsicum chinense) seeds were evaluated, with a view
toward determination of the best time for harvest. In
addition, enzymatic alterations were evaluated during the
germination process of the seeds processed at different
stages of maturity and storage.
MATERIALS AND METHODS
The research was conducted in the experimental area and in the
Central Seed Laboratory (Laboratório Central de Sementes) of the
Agriculture Department (Departamento de Agricultura) of the
Federal University of Lavras (Universidade Federal de Lavras -
UFLA), in Lavras, MG. The city located in the Southern Region of
Minas Gerais, latitude 21° 14’ S and longitude 40° 17’ W and at
918.8 m altitude. The annual average temperature is 19.4°C and
rainfall is distributed principally from October to April, with annual
amounts of 1529.7 mm. In a first stage of research, yellow
habanero pepper (C. chinense) seedlings were formed for
installation of the experiment in the field. Seeds were sown in
“styrofoam” trays with 72 cells containing the commercial substrate
Plantmax® - hortaliças and 5 ml of 2000 ppm solution of
ammonium sulfate per cell. Transplanting of the seedlings was
performed 45 days after seeding to an experimental area of the
olericulture sector of the Agriculture Department in an area with a
Dark Red Latosol/Oxisol and clayey texture; prepared
conventionally. Tests were installed in a randomized complete
block design with four replications. Each plot consisted of 2 rows of
5 m length with 5 plants per meter and spacing of 1.5 m between
rows. Plant cultivation was performed in accordance with Filgueira
(2005). Temperature and relative air humidity data during plant
development are presented in Table 1. Seeds were manually
extracted from many fruits at three stages of development: E1 (fruit
with first signs of yellowing), E2 (mature fruit) and E3 (mature fruit
submitted to seven days of rest). Then the seeds were dried in a
laboratory oven with air circulation at 35°C until reaching 8% water
content. The seeds corresponding to each stage of development of
the fruit were packed in airtight plastic packages and stored in a
walk-in cooler at 10°C and 50% relative humidity for periods of 0, 4
and 8 months after drying. At the end of each storage period, seed
quality was evaluated by means of germination tests; emergence
tests (Brazil, 2009); emergence rate index (Maguire, 1962);
electrical conductivity (Vidigal et al., 2008) and accelerated aging
(Bhering et al., 2006). Furthermore, the activity of the enzymes
esterase, superoxide dismutase (SOD), malate dehydrogenase
(MDH), alcohol dehydrogenase (ADH) and endo-β-mannanase
(Downie et al., 1994) was evaluated.
The activities of the enzymes α-amylase (Alfenas et al., 1991),
endo-β-mannanase, esterase, MDH and ADH were evaluated
during the germination process of the seeds at 0, 48, 96 and 144 h
after seeding. The seeds were ground in the presence of PVP
(polyvinylpyrrolidone) and liquid nitrogen in a mortar over ice and
afterwards stored at a temperature of -86°C. The experimental
design used was randomized complete blocks in a 3 × 3 factorial
design, with the factors being: stage of development of the fruit (E1,
E2 and E3) and storage periods (0, 4 and 8 months). Analysis of
variance was performed for all tests using the statistical program
Sisvar (Ferreira, 2000). For comparison among the means, the
Scott-Knott test was used at the 5% probability level.
RESULTS
It is observed from the results in Table 2, the interaction
between the factors stage of development of the fruit (E1,
E2 and E3) and storage periods (0, 4 and 8 months) was
significant by F test for all the variables analyzed. Lower
germination values were observed for the habanero
pepper seeds processed in the first stage of development
(E1), in all storage periods (Table 2). At 0 and 8 months
of storage, there was no difference between the
germination values of the seeds processed at the E2 and
E3 stages of development, whereas at 4 months of
storage, greater germination values were observed for E3
stage seeds. An increase in germination was observed
for E2 and E3 stage seeds at the 4th month of storage
Caixeta et al. 629
Table 2. Percentage of germinated seedlings (%), percentage of seedling emergence (%), emergence rate index and vigor
obtained by the accelerated aging test and electrical conductivity of habanero pepper seeds gathered at different stages of
development throughout the storage period.
Storage periods
Stages
0
4
8
Germination
E1
1Ba
3Ca
7Ba
E2
25Ac
41Bb
58Aa
E3
32Ab
50Aa
53Aa
Emergence
E1
5Cb
35Ba
30Ba
E2
60Bb
85Aa
84Aa
E3
75Ab
86Aa
87Aa
Emergence rate index
E1
0Cb
5Ba
5Ca
E2
8Bc
23Ab
29 Ba
E3
13Ac
25Ab
34Aa
Accelerated aging
E1
1Cb
3Cb
23Ca
E2
20Bc
68Ab
87Ba
E3
43Ab
47Bb
95Aa
Electrical conductivity
E1
825Cb
580Bb
634Ca
E2
750Bc
535Ab
463Ba
E3
654Ac
511Ab
406Aa
(1) Means followed by the same capital letter in the column and small letter in the row do not differ among themselves by the Scott-
Knott test at the 5% probability level.
(Table 2). These values were maintained in E3 stage
seeds but increased in E2 stage seeds after eight months
of storage. As for E1 stage seeds percentage of
germinated seedlings were very low and did not differ
significantly during storage. The results obtained in the
emergence test and emergence rate index, under
greenhouse conditions, are presented in Table 2. Lower
plantlet emergence values were observed in seeds
processed in the E1 development stage at all storage
periods. At 4 months of storage, there was an increase in
the percentage of seedling emergence from seeds
processed in different stages of development (Table 2).
The results of the emergence rate index indicated less
vigor in the E1 stage seeds and greater vigor values in
the E2 and E3 stage seeds stored for 4 months.
However, at 8 months, there was vigor increase for E3
and E2 seeds and maintenance for E1 stage seeds. The
results of the accelerated aging test (Table 2) showed
greater means of seed vigor in E1, E2, and E3 stage
seeds stored for 8 months. At the beginning of storage,
the seeds processed in the E3 stage were more vigorous
for other stages. Regardless of the storage period, less
vigor was observed in habanero seeds extracted at the
E1 stage. By the electrical conductivity test (Table 2),
greater vigor was observed in E3 seeds that were either
recently stored, or stored for 8 months. There was no
significant difference in the conductivity values observed
in E2 and E3 stages seeds stored for 4 months. In the
same way, greater leaching was also observed at 8
months of storage (Table 2). Regardless of the stage of
development of habanero pepper seeds, at the end of 8
months of storage, there were lower means of electrical
conductivity.
Considering the electrophoretic analyses, the
enzymatic profile of esterase (Figure 1) +showed a
greater activity of this enzyme for E1 stage seeds at the
three storage periods, with this being attributed to greater
immaturity of these seeds but also to seed quality
deterioration throughout the storage period. Regarding
enzyme SOD (Figure 1), there was an increase in its
630 Afr. J. Agric. Res.
P0
P4
P8
1 2 3 1 2 3 1 2 3
ESTERASE
B
1 2 3 1 2 3 1 2 3
P0
P4
P8
SOD
ADH
MDH
1 2 3 1 2 3 1 2 3
P0
P4
P8
1 2 3 1 2 3 1 2 3
P4
P0
P5
Figure 1. Esterase enzyme profiles, SOD, MDH and ADH of habanero pepper seeds processed in the E1 (1), E2 (2) and E3 (3)
stages in the three storage periods 0 (P0), 4 (P4) and 8 (P8) months of storage.
activity at 4 months of storage for all the three stages of
development. At 8 months of storage, less activity was
observed in E2 and E3 seeds, while in E1 stage seeds
there was an increase in the activity of this enzyme. The
enzyme MDH (Figura 1) had increased activity in all three
stages of development at 8 months of storage. Regarding
the enzyme ADH (Figure 1), greater activity in E2 stage
seeds was observed at 4 and 8 months. The profile of the
enzyme endo-β-mannanase showed an increase in
activity (Figure 2) for seeds processed in the most
advanced stages of development. Moreover, less activity
of this enzyme was observed in recently stored seeds
regardless of the maturity stage. In the present research,
greater germination and vigor values were observed after
storage of the seeds in E2 and E3 stage, leading to the
supposition of breaking of dormancy of the seeds during
storage (Table 2). Considering the electrophoretic
analyses of the enzymatic patterns during germination,
there was no difference in the activity of the enzymes in
the dry seeds in relation to those soaked for 48 h in all
the enzymatic patterns, with the exception of that
observed for the enzyme endo-β-mannanase (Figure 3).
Variations in the MDH enzyme patterns in habanero
pepper seeds (Figure 4) were verified, with less activity in
the period of 144 h of soaking in seeds processed in the
different stages of development and different storage
times. In relation to the patterns observed for the enzyme
ADH, it may be observed that with the advance of the
soaking period, the activity of the enzyme ADH
diminished in the three storage periods (Figure 4). It was
also verified that activity of the enzyme increased
throughout the storage period in seeds soaked for 48 h.
Like MDH, there was variation in the activity of the
enzyme ADH in seeds processed at different stages of
maturity, in terms of the storage period and of the
soaking period. In habanero seeds (Figure 4), greater
activity of the ADH enzyme was observed in E2 and E3
seeds after 48 h of soaking at 0 and 8 months of storage,
whereas at 4 months of storage, greater activity was
observed in seeds of all three stages of development
after 48 h of soaking.
Regarding the enzyme α-amylase, Figure 4 showed
that there was variation in the activity of this enzyme in
terms of the stage of development at which the seeds
were processed, storage period and duration of soaking.
The activity of the enzyme α-amylase may become
evident through the clearer bands in a bluish background,
where the starch was hydrolyzed. Greater activity of the
enzyme α-amylase was observed at 4 months of storage
compared to 0 and 8 months of storage for all stages of
Caixeta et al. 631
0
50
100
150
200
250
300
E1 - 0
E1 - 4
E1 - 8
E2 - 0
E2 - 4
E2 - 8
E3 - 0
E3 - 4
E3 - 8
Atividade (picomol.min-1.g.-1)
Atividade (picomol.min-1.g.-1)
Atividade (picomol.min-1.g.-1)
Figure 2. Activity of the enzyme endo-β-mannanase in habanero pepper seeds processed in the E1, E2 and E3
development stages in three storage periods of 0, 4 and 8 months of storage.
seed development. E1 seeds presented greater activity of
this enzyme than E2 and E3 seeds at the different
soaking times and storage periods, except for recently
stored seeds soaked for 48 h and those stored for 8
months and soaked for 96 h.
DISCUSSION
Barbedo et al. (1994) also verified that better quality
eggplant seeds were obtained from fruit harvested 50
days after anthesis and submitted to 15 days of post
harvest storage. According to Sanchez et al. (1993),
green pepper seeds should remain in the mature fruit (50
days after anthesis) after harvest from 7 to 14 days so
that maximum germination potential is reached.
According to Nascimento et al. (2006), immature fruit, of
green color, generally produces seeds with low vigor and
germinating power or even poorly formed seeds. The low
germination percentage at the beginning of storage may
be related to the presence of dormancy in the seeds
which was broken throughout the storage period. These
results corroborate those found by Bosland and Votava
(1999), in which dormancy was observed in recently
gathered seeds of species of the genus Capsicum (Table
2). Thus, it may be inferred that in recently stored seeds
and in those stored for 4 months, aerobic respiration is
greater at the beginning of the germination process. It
may be observed that the greatest percentage of
germination is after 4 months of storage. According to
Nascimento et al. (2006), the sowing of pepper seeds
recently extracted from the fruit may represent a risk for
obtaining uniform stands, contributing to increased seed
expenses. These seeds are induced dormancy to
preserve the perpetuation of the species. Since this seed
peppers should be stored before being sown. The
increase in seedling test emergency seed processed in
different stages of development in relation to germination
emphasizes that in spite of the reports regarding the
occurrence of dormancy in pepper seeds (Bosland and
Votava, 1999) (Table 2). This dormancy can be broken
down by microorganisms in the substrate, when the
seeds of determined cultivars are extracted from
completely mature fruit and seeded thereafter (Bolsland
and Votava, 1999).
Randle and Honma (1981) verified in work with
different cultivars of the genus Capsicum, that the
genotype and the age of the fruit influence the intensity of
dormancy of the seeds. The authors reported that seeds
extracted from mature fruit with days of rest before seed
extraction germinate more rapidly, younger fruits being
more prompt to increased seed dormancy. According to
Barbedo et al. (1994), by the emergence rate index, it is
possible to detect small existing differences in the
physiological quality of cucumber seeds extracted from
fruits harvested 15 to 45 days after anthesis (DAA) and
without storage. Valdes and Gray (1998), upon
harvesting tomato fruits of differing maturity stages and
without post harvest storage, observed that the mean
germination time of the seeds differed significantly among
632 Afr. J. Agric. Res.
0
50
100
150
200
250
300
350
E1(0-0h)
E1(0-48h)
E1(0-96h)
E1(0-144h)
E1(4-0h)
E1(4-48h)
E1(4-96h)
E1(4-144h)
E1(8-0h)
E1(8-48h)
E1(8-96h)
E1(8-144h)
Atividade (picomol.min-1.g.-1)
0
50
100
150
200
250
300
350
E2(0-0h)
E2(0-48h)
E2(0-96h)
E2(0-144h)
E2(4-0h)
E2(4-48h)
E2(4-96h)
E2(4-144h)
E2(8-0h)
E2(8-48h)
E2(8-96h)
E2(8-144h)
Atividade (picomol.min-1.g.-1)
0
50
100
150
200
250
300
350
E3(0-0h)
E3(0-48h)
E3(0-96h)
E3(0-144h)
E3(4-0h)
E3(4-48h)
E3(4-96h)
E3(4-144h)
E3(8-0h)
E3(8-48h)
E3(8-96h)
E3(8-144h)
Atividade (picomol.min-1.g.-1)
E1
E2
E3
Atividade (picomol.min-1.g.-1)
Atividade (picomol.min-1.g.-1)
Atividade (picomol.min-1.g.-1)
Figure 3. Activity of the enzyme endo-β-mannanase in habanero pepper seeds during 0,
48, 96 and 144 h of germination in the E1, E2 and E3 stage in the three storage periods of
0, 4 and 8 months of storage.
maturity stages, greater germination time being observed
in the less mature seeds. Our findings on electrical
conductivity were similar to those observed by Vidigal et
al. (2006) and Dias et al. (2006) for tomato seeds
extracted from fruits with different stages of maturity
submitted to post-harvest storage (Table 2). Greater
leaching of exudates was observed in immature seeds
consistent with less structuring of the organelle and
cellular membrane system. Nevertheless, the high
conductivity value observed for E1 seeds suggests
destructuring of the system of membranes, probably
because of their immaturity (Albuquerque et al., 2009),
and this fact is also reinforced by the results of the other
physiological tests.
As observed by Bhering et al. (2006), the accelerated
aging test was efficient to evaluate the effect of pepper
seeds, checking statistical difference between the
different treatments (Table 2). Esterase is an enzyme that
participates in membrane hydrolysis of esters. This fact
shows greater lipid peroxidation since this enzyme is
involved in ester hydrolysis reactions, being directly
connected to lipid metabolism (Santos et al., 2004). Many
of these lipids are constituents of membranes, whose
degradation increases with deterioration (Figure 1). The
enzyme endo-β-mannanase being involved in the
degradation of the endosperm in seed germination. In
lettuce and coffee seeds, this enzyme is considered as
key in the germination process, being involved in
mannanase degradation at the time of germination,
resulting in weakening of the cell walls of the endosperm
(Silva et al., 2004; Veiga, 2005) (Figure 2). Vidigal et al.
(2009) observed a small increase in the activity of SOD in
chili peppers obtained from fruits harvested 50 DAA and
stored for 6 days, greater physiological quality as
evaluated by the germination and vigor tests (Figure 1).
Enzyme ADH activity reported here was also found by
Vidigal et al. (2009) (Figure 1). This enzyme is related to
anaerobic respiration, promoting reduction of the
acetaldehyde to ethanol. Acetaldehyde accelerates seed
deterioration (Buchanan et al., 2005). With the increase
of ADH activity, the seeds are more protected against the
deleterious action of this compound, which is greater
when compared to that of ethanol. In the 8th month of
storage, through the fact of the seeds being in more
advanced stages of deterioration, there is greater
respiratory intensity and consequently greater demand of
activity from the enzyme MDH (Figure 1). MDH enzyme
patterns varied according to storage and soaking periods
and seed maturity stages. In research undertaken by Taiz
and Zeiger (2004), no difference in MDH activity was
observed in seeds during the maturation process.
Nevertheless, the authors reported that the reserve
organs in development need greater energy supply and,
therefore, respiratory activity in these plant tissues is
more intense (Figure 4). These results with MDH and
ADH may be associated with the germination and vigor
data, in which the seeds processed in the E2 and E3
Caixeta et al. 633
stages have better quality than the seeds of the E1 stage
(Table 2).
According to Nedel et al. (1996), within a group of
enzymes, the α and β - amylases are involved in the main
starch degradation system. Development of the amylase
activity constitutes an important event and may be
detected at the beginning of germination with its main
role being the making of substrates for the plantlet
nutrition until it becomes photosynthetically self-sufficient.
A large number of types of seed dormancy arise from
blocking the action of α-amylase. The α-amylase present
in the dormant seeds is found in small quantities. The
activity of this enzyme increases to the extent that the
dormancy of rice seeds is overcome during the storage
period (Vieira et al., 2008).
In this study, the presence of dormant seeds was
observed, principally at the E1 stage of development and
in recently stored seeds (Table 2). In these seeds, high
activity of α-amylase was observed, which confirms the
importance of this enzyme in the germination process of
pepper seeds. In E1, E2 and E3 stage seeds and
recently stored seeds, there was an increase in the
activity of the enzyme endo-β-mannanase (Figure 3) to
the extent that the soaking period of the seeds was
increased during the germination process. The greatest
activity of this enzyme was observed at 144 h of soaking,
which coincided with the occurrence of root protrusion.
Seeds of the three stages of development stored for 4
and 8 months germinated 96 h after soaking. These data
may be correlated with the greater activity of this enzyme
in seeds submitted to these treatments. Greater activity
of the endo-β-mannanase, in absolute values, was
verified in the E2 and E3 stage seeds at all the storage
and soaking periods during the germination process. The
lowest germination and vigor values were observed in E1
seeds and in recently stored seeds. In these seeds, the
activity of the enzyme endo-β-mannanase was, the
lowest, indicating the importance of this enzyme in the
germination of pepper seeds. The greatest level of
activity of this enzyme was observed in seeds stored for
4 months and soaked for 96 h, and this was true for all
the three stages of development (Figure 3). The results of
the tests used for evaluation of physiological quality,
showed an increase in the germination and vigor values
for 4 month-stored seeds (Table 2). Based on these
results, the presence of dormancy in recently harvested
seeds is inferred. This dormancy has probably been
overcome in the 4th month of storage as shown by a
greater activity of the enzyme endo-β-mannanase.
Conclusions
By means of the physiological tests and the activity of the
enzymes evaluated, it was observed that habanero
pepper seeds should be extracted from E3 stage fruits,
which ensures production of better quality seeds. Seed
634 Afr. J. Agric. Res.
P0
0 48 96 144
1 2 3 1 2 3 1 2 3 1 2 3
P4
1 2 3 1 2 3 1 2 3 1 2 3
1 2 3 1 2 3 1 2 3 1 2 3
P8
P8
MDH
0 48 96 144
0 48 96 144
P0
1 2 3 1 2 3 1 2 3 1 2 3
P4
1 2 3 1 2 3 1 2 3 1 2 3
1 2 3 1 2 3 1 2 3 1 2 3
0 48 96 144
0 48 96 144
0 48 96 144
ADH
P0
1 2 3 1 2 3 1 2 3 1 2 3
P4
1 2 3 1 2 3 1 2 3 1 2 3
1 2 3 1 2 3 1 2 3 1 2 3
P8
P8
0 48 96 144
0 48 96 144
0 48 96 144
ESTERASE
1 2 3 1 2 3 1 2 3 1 2 3
P0
P4
1 2 3 1 2 3 1 2 3 1 2 3
1 2 3 1 2 3 1 2 3 1 2 3
α-AMILASE
0 48 96 144
0 48 96 144
0 48 96 144
Figure 4. Electrophoretic patterns of the enzyme MDH, ADH, esterase and α-amylase observed in habanero pepper seeds during
germination: 0, 48, 96 and 144 h in the E1 (1), E2 (2) and E3 (3) stages with 0 (P0), 4 (P4) and 8 (P8) months of storage.
extraction from fruits harvested at E1 stage must be
avoided because of lower physiological quality due to
seed dormancy and immaturity. Seed physiological
maturity in the species studied does not coincide with the
maximum germination and vigor values due to the
incidence of dormancy.
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