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Physiological and biochemical alterations during germination and storage of habanero pepper seeds

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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.
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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)
Tm (ºC)
UR (%)
Pt (mm)
I (h)
January
27.6
21.9
85.7
554.7
3.1
February
28.9
22.6
73.1
151.3
7.5
March
30.9
23.6
66.5
35.4
9.0
April
28.1
21.8
72.4
35.6
7.3
May
25.7
18.1
70.6
30.4
7.5
June
25.8
17.3
66.3
5.9
8.8
July
25.4
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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... For the DWTS, higher values were obtained after post-harvest resting. Similar results were reported in other pepper seeds [23][24][25]. ...
... Studies of the sweet pepper cultivar Amarela Comprida demonstrated increases in DWTS up to 75 DAA, with a maximum weight of 6.6 g [28]. However, in the pepper cultivar 'Malagueta', the maximum DWTS obtained was 3.2 g at 80 DAA, and in the 'Biquinho' pepper, it was 2.5 g at 70 DAA [25], thus showing that values can vary according to genotype. ...
Article
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There are few studies about the influence of fruit maturation and post-harvest resting on seed composition, which can be necessary for seedling development and future establishment. Thus, the objective of this study was to evaluate the effect of maturation and post-harvest resting of fruits on the macronutrient and protein content of sweet pepper seeds. The experimental design was a randomized block, with eight treatments, in a 4 × 2 factorial arrangement. The first factor was fruit maturation stages (35, 50, 65 and 80 days after anthesis), and the second, with and without post-harvest resting of the fruits for 7 days. The characteristics evaluated in seeds were the dry weight of one thousand seeds, macronutrient content, and content of albumin, globulin, prolamin and glutelin proteins. There were reductions in K, Ca and Mg content, and an increase in seed content of albumin, globulin and prolamins as a function of the fruit maturation stage. Post-harvest resting of the fruits provided higher Ca content and protein albumin in seeds. The decreasing order of macronutrients and protein content in seeds, independent of fruit maturation and resting stage of the fruits, was N > K > P > Mg > S > Ca, and albumin > globulin ≈ glutelin > prolamine, respectively.
... In 'Amarela Comprida' pepper seeds, at 40 and 50 DAA, there was no difference in the enzymatic response, which could be associated with the maturation stage, however, there was a slight increase in SOD activity in seeds obtained from fruits harvested from 50 DAA and stored for 6 days [9]. This enzyme, in 'Habanero' pepper seeds, showed higher and lower enzyme activity when fruits that were harvested with the first signs of yellowing and completely ripe, respectively [30]. ...
Article
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Abstract The post-harvest resting of the fruits can improve seed physiological quality ,once it allows the seed to complete the maturation process, so it has been a common practice in vegetable seed companies, however, there are a few studies of this technique in sweet pepper. The objective of this research was to evaluate physiological quality, and biochemical response of sweet’s peppers in regarding on the stage of maturation and the post-harvest rest of the fruits. The experimental was conducted in a 4x2 factorial, being the first factor comprised four maturation stages (35, 50, 65 and 80 days after anthesis) and, the second the post-harvest management of fruits, with and without a temporary storage of seven days. Seeds were evaluated for water content, weight of thousand seeds, germination, vigor, superoxide dismutase, catalase and peroxidase activities, lipid peroxidation and hydrogen peroxide content. Fruit harvest time indicated is 80 days after anthesis (fruits 100% yellow) when seeds showed maximum germination and vigor. The post-harvest resting of the fruits was beneficial to seed physiological quality, weight of one thousand seeds and to reduce hydrogen peroxide content. Seeds of higher physiological quality showed lower superoxide dismutase and catalase enzyme activity, so they can be used as a marker of physiological quality in sweet pepper seeds.
... There are numerous studies on longevity of conventionally-produced pepper seeds (Thanos et al., 1989;Sundstrom, 1990;Sanchez et al., 1993;Basay et al., 2006;Demir and Ozcoban, 2007;Demir et al., 2009;Caixeta et al., 2014;Panayotov and Aladjadjiyan, 2014;Adam et al., 2018;Verma et al., 2018). Nevertheless, organic seed production systems are different than conventional ones and require specific measures for production (Anonymous, 2010). ...
Article
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This study was carried out to determine seed longevity in organic and conventionally produced pepper seeds from four different pepper cultivars. Seeds were stored at 20± 2 o C with 7.5±0.5 seed moisture over 48 months. Longevity of seeds were evaluated with Ki (initial seed quality), P50 (half-viability period), σ (standard deviation of distribution of seed deaths in time), and regression coefficient values. The highest longevity was observed in 'Corbaci' and 'Yaglik' cultivars, while 'Surmeli' and 'K. Dolma' were found to have shorter longevity. P50 was 43.4 and 40.2 months for 'Corbacı' and 34.9 and 39.7 months for 'Yaglık' organic and conventional cultivars, respectively, whereas it was about 21.4 and 23.7 months in 'K. Dolma' and 'Surmeli' cultivars. Similarly, the highest σ and regression coefficient values were observed for 'Corbaci' and the lowest for 'Surmeli' cultivars. Organic and conventional pepper seed longevity was not different in the same species. Regression coefficient values were 0.043 in organic and 0.046 in conventional seeds for 'Corbaci'. Very close values were found between the two production systems for the other cultivars too. Results indicate that organic seeds had similar longevity to conventional ones. The main differences originated from the cultivars, not from the production system.
... A cultura atualmente deixa de ser considerada de segundo plano e passa a assumir maior importância na produção hortícola no país, isso ocorre devido a gama de produtos que possuem a pimenta (Capsicum frutescens L.) como matéria-prima, sendo comercializada In natura, conserva, páprica, pastas, geleias, molhos, em pó, desidratadas e ornamentais agregando maior valor ao produto contribuindo para a ampliação do setor (CAIXETA et al. 2014). Filgueira (2008) afirma que o aumento no consumo de pimentas deve-se a estudos que comprovaram que pimentas apresentam teores de vitamina A e C superiores aos encontrados no pimentão e demais olerícolas produzidas no Brasil. ...
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Objetivou-se avaliar modelos de evapotranspiração de referência e estimar os coeficientes de cultivo da pimenta. O experimento foi conduzido no período de 27/04 a 09/09/2015, na área experimental do Grupo Irriga do Campus Arapiraca da UFAL. Para encontrar o consumo hídrico da cultura da pimenta foi calculado o Kc da pimenta pela razão da ETc da pimenta pela ETo (estimada pelos métodos de Penman-Monteith , Hargreaves-Samani, Radiação-Solar, Blaney-Criddle e de Priestley-Taylor). Para comparação e análise dos métodos de ETo’s foram utilizados critérios envolvendo o erro padrão de estimativa (EPE), erro padrão da estimativa ajustado (EPEa), coeficientes de correlação (r) de Pearson e de determinação (r2), índice de desempenho (c) e do índice de concordância (d). Para as análises de correlação e de regressão linear entre os métodos de estimativas de ETo e os gráficos foi utilizado o software estatístico RStudio®. Os resultados mostraram que o consumo hídrico diário da cultura da pimenta foi de 4,32 mm dia-1. O método de Radiação Solar apresentou o melhor desempenho na estimativa da ETo em relação ao método padrão-FAO Penman-Monteith. O Kc da pimenta, recomendado para a região semiárida, é 0,70; 1,16; 1,23 e 1,00, para as fases I, II, III e IV respectivamente, obtidos pelo método padrão-FAO. Na ausência de informações agrometeorológicas para se estimar a ETo pelo método padrão-FAO, recomendam-se os valores de Kc de 0,81; 1,22; 1,28 e 0,98, obtidos pelo método de ETo de Radiação-Solar. Também poderá usar a equação de ajuste obtida pela correlação entre as EToRS e EToPM, possibilitando o uso do KcPM.
... The region where the radicle emerges, called endosperm cap, is composed of high cellulose and hemicellulose polysaccharide content, which confers resistance and protection to the embryo (SILVA et al., 2005;EIRA et al., 2006). Some authors have affirmed that for seed germination to occur, the endosperm cap must be weakened through the activity of the endo-β-mannanase enzyme, as occurs in seeds of several species, such as Datura ferox (ARANA et al., 2006), peppers (Capsicum spp.) (CAIXETA et al., 2014), coffee (Coffea arabica L.) (SILVA et al., 2004), and lettuce (Lactuca sativa) (ALBUQUERQUE et al., 2010). ...
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Enzymes play a fundamental role in degradation of molecules during seed germination, development, and deterioration. Endo-β-mannanase is one of the main enzymes responsible for hydrolysis of mannans in the endosperm during germination of coffee seeds through its action in hydrolytic degradation of cell walls and in weakening the structures of the endosperm that surround the embryo, allowing radicle emergence. The aim of this study was to determine the activity of the endo-β-mannanase enzyme in the structures of coffee seeds for the purpose of assessing the relationship between this activity and the physiological quality of the seeds under different processing and drying methods. Coffea arabica L. fruit in the cherry maturity stage was subjected to three different types of processing: natural (seeds maintained in the fruit itself), fully washed (fruit pulped mechanically and the seeds demucilaged by fermentation in water), and semi-washed or demucilaged (both fruit pulp and mucilage removed mechanically); and two methods of drying: slow drying (suspended screen) in the shade, and rapid drying in mechanical dryer at 35°C to a moisture content of 11±1%. After processing and drying, the physiological quality of the seeds was evaluated through the germination test, and endo-β-mannanase enzyme activity was quantified. Coffee seeds submitted to natural processing have lower physiological performance, as well as greater deterioration and greater activity of the endo-ß-mannanase enzyme. Removal of mucilage during fully washed and semi-washed processing of coffee seeds reduces the activity of the endo-ß-mannanase enzyme and lowers deterioration, especially after faster drying. The enzyme endo-ß-mannanase is efficient in studying of the effects of processing and drying on coffee seeds, and can be evaluated in whole seeds, endosperms or embryos.
... First germination count and seedling emergence also increased after four months, but there was reduction after eight months of storage. Increment in pepper seed germination during storage is an indication of breaking of secondary dormancy ( Caixeta et al., 2014;Lima & Smiderle, 2014). The accelerated aging test was proposed to evaluate seed storage potential ( Marcos Filho, 2015). ...
Article
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Drying air temperature has direct influence on seed quality, especially for vegetable species harvested with high moisture content. This study aimed to determine and model the drying curves of ‘Cabacinha’ pepper seeds (Capsicum chinense L.) at different temperatures, and to evaluate their physiological quality after drying and during storage. The experimental design was completely randomized in a 3 x 3 factorial scheme, with three drying temperatures (35, 38 and 42 °C) and three storage periods (0, 4 and 8 months), with four replicates. Seed quality was evaluated by germination test, first germination count, accelerated aging, electrical conductivity, seedling emergence, and speed of emergence. The model Two Terms is that best represents the drying curves of ‘Cabacinha’ pepper seeds at temperatures of 35, 38 and 42 °C. Higher germination and vigor values are obtained in seeds dried at 35 and 38 °C. Drying at 42 °C and storage reduce the physiological quality of the seeds, regardless of the drying temperature studied.
... In interpreting the results of the height of the tomato plants, no significant difference between the analyzed treatments were observed (Fig. 1), although the bioprotector application in all rates showed greater responses. Similar results were obtained for green peppers in field conditions, improving the effects of the fertilization treatments (Santana et al., 2014) and by Caixeta et al. (2014) during germination. The control treatment (earthworm compost 20 t ha -1 ) presented low values and were inferior to the bioprotector, particularly at higher rates, and the soluble fertilizer at the recommended rate. ...
Article
The production of biofertilizers from rocks is an economic process that increases nutrients for plant nutrition and reduces environmental pollution. The aim of the study was to evaluate the effectiveness of the bioprotector from phosphate and potassium rocks mixed with an earthworm compound inoculated with free-living diazotrophic bacteria (Beijerinckia indica) and fungus containing chitosan (Cunninghamella elegans) on the characteristics of tomatoes, peppers and green peppers inoculated and non-inoculated with the wilt bacteria Ralstonia solanacearum. The fertilization treatments were: bioprotector applied at the 50% recommended rate - RR (NPKP50), 100% RR (NPKP100) and 150% RR (NPKP150); soluble fertilizers applied alone at the recommended rate RR (NPKF100), with antibiotic (NPKF100+ant) and with crustaceous chitosan (NPKF100+C). A control treatment with earthworm compound (20 t ha⁻¹) was used for comparative purposes. The best results for all horticultural plants were obtained with the bioprotector (NPKP) applied at the highest rate (150% RR) and with the soluble fertilizer (NPKF) at the recommended rate for plants not inoculated with Ralstonia solanacearum. With regard to the effect of the bioprotector in the control of Ralstonia solanacearum, a different response for the horticultural crops with peppers revealed normal growth when inoculated with the pathogenic bacteria, and all of the tomato plants with soluble fertilizer alone died after pathogen addition. Soluble fertilizer with the addition of antibiotic and crustaceous chitosan protected green peppers and peppers against R. solanacearum, particularly when using chitosan. The bioprotector (NPKP) appears to be an alternative to soluble fertilizer.
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Objetivou-se avaliar a produção, rendimento, qualidade e biometria de frutos de pimenta não pungentes cultivados em ambiente protegido por meio da poda de hastes caulinares e raleio dos frutos. Os tratamentos foram compostos por sistemas de condução das hastes (sem poda e com poda) e raleio (sem raleio e com raleio dos frutos). O delineamento experimental foi blocos casualizados com quatro repetições em esquema fatorial 2 (sem poda e com poda das hastes) x 2 (sem raleio e com raleio dos frutos). Foram avaliados os frutos de quatro plantas por parcela, totalizando oito momentos de colheita. A poda promoveu incremento na fitomassa de frutos totais e comerciais, número de frutos comerciais, comprimento (C), diâmetro (D), relação C/D, sólidos solúveis totáis, pH e acidez total titulável dos frutos de pimenta. O raleio de frutos promoveu incremento do comprimento e diâmetro médio dos frutos de pimenta. Recomenda-se a poda das hastes das plantas de pimenteira para obter maior massa média e diâmetro de frutos.
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Apesar de parecer pouco importante para a agricultura, o setor produtivo de pimentas (Capsicum) é muito extenso e promissor, fazendo parte fundamental do agronegócio brasileiro. O objetivo desse trabalho foi realizar uma revisão narrativa apontando os principais fatores referentes a utilização de diferentes métodos de secagem e temperaturas de armazenamento em sementes de pimenta, e como afetam a manutenção da qualidade fisiológica. Quanto à metodologia, utilizou-se o modelo de estudo baseado em pesquisa bibliográfica exploratória e abordagem qualitativa, trazendo informações a respeito da secagem e armazenamento adequados na manutenção da qualidade de sementes de pimenta, abordando os aspectos e fatores que interferem diretamente nesses processos. Observou-se através da literatura consultada que a secagem e armazenamento influem diretamente na qualidade das sementes, mas se faz importante destacar que processos no campo, como a época adequada de colheita, também impactam significativamente. Como o setor está em desenvolvimento, há uma necessidade significativa de pesquisa em tecnologia de produção de sementes com alta qualidade genética, física, fisiológica e de sanidade que resultem maior produtividade final, sendo a secagem e o armazenamento etapas cruciais desse processo. Através dos achados foi possível observar o quão importante é a realização da secagem e armazenamento adequados para manutenção do vigor, utilizando temperaturas e condições especificas conforme exigido para as sementes de pimenta. Embora seja elevado o número de publicações sobre o manejo e produção de pimentas, são necessários estudos mais específicos que identifiquem a otimização do processo produtivo de sementes, bem como as operações de secagem e armazenamento.
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The appropriate fruit harvest stage for optimum seed quality in four pepper genotypes usually grown in Minna, Niger State was determined. Mass transplanting of pepper seedlings was carried out on beds at 50 and 75 cm intra and inter row spacings respectively. Flowers were date-tagged as they opened for about 40 days. Fruits of different ages (24, 28, 32, 36, 40, 44, 48 and 52 DAA) were harvested at once. Data were taken on fruit colour, seed dry matter, germination percentage (GP), germination rate index (GRI) and germination index (GI). Data collected were subjected to analysis of variance (ANOVA) using Minitab 17.0 version; means were separated using Tukey's Honest Significant Different Test. Fruit colour remained green up till 28 DAA. At 32 DAA, red colouration became noticeable in a small proportion of the fruit. This continued gradually until fruits became completely red outside at 52 DAA. No significant differences were recorded among the dry matter of seeds of the four genotypes while its values increase significantly from 19.67% to 64.03% from 24 to 52 DAA respectively. Seeds of 'Shombo' genotypes (V4) is superior in both viability and longevity than all the 'Tatashe' genotypes (V 1, 2 and 3); genotype V2 of 'Tatashe' however maintain viability longer than V 1 and 3. GP increased significantly from 8.00% to 79.25% from seeds harvested at 28 to 52 DAA respectively prior to storage with slight increase to 81.50% at 52 DAA when seeds were 14 days in storage. The trends recorded in GRI and GI were similar to that of GP. The general decreases in the values of GP, GRI and GI when seeds were beyond 14 days in storage was an indication that deterioration has set in. It is concluded from this study that pepper seeds reached maximum quality at 52 DAA when fruits were completely red outside.
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O momento da colheita e os métodos de secagem podem influenciar a qualidade das sementes de cafeeiro durante o armazenamento. Este trabalho foi realizado com o objetivo de avaliar os efeitos do estádio de maturação e do método de secagem sobre a qualidade fisiológica e a armazenabilidade de sementes de cafeeiro. Os ensaios foram realizados nos Laboratórios de Análise de Sementes e de Técnicas Moleculares do Departamento de Agricultura da UFLA. Os frutos do cultivar Rubi foram colhidos, despolpados, e as sementes lavadas e deixadas sobre papel para retirada da água superficial. As sementes nos estádios verde cana e cereja foram submetidas à secagem convencional (à sombra) e à secagem em secador estacionário sob temperatura de 35ºC. Como testemunha foram analisadas sementes sem secagem. As avaliações foram feitas imediatamente após os tratamentos de secagem e após 4 e 8 meses de armazenamento. As sementes foram armazenadas a 10ºC em sacos plásticos impermeáveis. Foram realizados os testes de germinação, de protrusão radicular, de matéria seca de plântulas, de índice de velocidade de emergência, de condutividade elétrica, além de análises eletroforéticas de proteínas resistentes ao calor e da quantificação da atividade da enzima endo-ß-mananase. O delineamento foi inteiramente ao acaso em esquema fatorial dois (estádios de maturação) x três (métodos de secagem) x três (tempos de armazenamento), com quatro repetições. Sementes de cafeeiro colhidas no estádio cereja têm maior potencial de armazenamento que no estádio verde cana. Ocorre redução de germinação e vigor nas sementes de cafeeiro colhidas no estádio verde cana, quando submetidas à secagem rápida. A presença ou intensidade de bandas de proteínas resistentes ao calor está associada à secagem das sementes. Ocorre maior atividade da enzima endo-ß-mananase em sementes colhidas no estádio cereja que no estádio verde-cana. Ocorre aumento da atividade da enzima endo-ß-mananase durante o armazenamento.
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Although thought of as a minor crop, peppers are a major world commodity due to their great versatility. They are used not only as vegetables in their own right but also as flavourings in food products, pharmaceuticals and cosmetics. Aimed at advanced students and growers, this second edition expands upon topics covered in the first, such as the plant's history, genetics, production, diseases and pests, and brings the text up to date with current research and understanding of this genus. New material includes an expansion of marker-assisted breeding to cover the different types of markers available, new directions and trends in the industry, the loss of germplasm and access to it, and the long term preservation of Capsicum resources worldwide.
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Seeds are uniquely equipped to survive as viable regenerative organisms until the time and place are right for the beginning of a new generation; however, like any other form of life, they cannot retain their viability indefinitely and eventually deteriorate and die. Fortunately, neither nature nor agricultural practice ordinarily requires seeds to survive longer than the next growing season, though seeds of most species are able to survive much longer under the proper conditions.
Article
Some peppers (Capsicum sp.) exhibit a primary seed dormancy at harvest. This study was designed to examine the effect of fruit maturity, dry and moist seed storage and seed postharvest maturation treatments on the seed germination of four bell pepper cultivars (Capsicum annuum L., cultivars ‘Early Calwonder’, ‘Resistant Giant No. 4’, ‘VR2’ and ‘Yolo Wonder’). In 1987 and 1988, fruits were harvested 30 (mature green), 40 (breaker), 50 (mature red) and 60 (over-mature red) days postanthesis (dpa), and seeds were extracted for dry storage treatments, or allowed to remain in fruits for postharvest maturation treatments of 0, 7, 14, 21 and 28 day periods. In 1988, seeds were also stored over water or saturated salt solutions which provided relative humidities of 7, 51 and 97.5% to determine the influence of seed moisture during seed storage. Seeds from red and over-mature red (50 dpa and 60 dpa, respectively) fruit generally had greater dry weights and higher germination percentages relative to seeds from less mature fruits. Seeds extracted from mature green (30 dpa) fruit did not germinate regardless of storage time. Seed postharvest maturation of up to 14 days in green fruit significantly increased seed germination in all cultivars. When pepper seeds were allowed to remain in mature red (50 dpa) fruit for various postharvest maturation periods, germination percentages were generally significantly greater than in dry storage and mature green (30 dpa) postharvest maturation treatments. Starch concentrations of ‘Resistant Giant No. 4’ seeds increased from 10 to 80 mg glucose per 100 g dry wt. when seeds remained in fruit for 4 weeks after harvest. Dry storage of seeds following extraction from red fruit did not significantly increase germination, indicating that seed afterripening did not occur. Short storage periods (7 and 14 days) at low relative humidity (7 and 51%) significantly improved seed germination of ‘Early Calwonder’ and ‘Resistant Giant No. 4’. Storage for 7–14 days of ‘Early Calwonder’ and ‘Resistant Giant No. 4’ seeds at high (97 and 100%) relative humidities was also beneficial, but at longer periods germination percentages of all cultivars dropped. There were few significant differences in seed dry weight between dry storage and postharvest maturation treatments. It is possible that in situ priming may have occurred in 30 and 50 dpa seed postharvest maturation treatments. These results suggest that following fruit harvest, pepper seeds should remain in red (50 dpa) fruit for a short postharvest maturation period to achieve maximum seed germination potential.
Article
Studies were conducted on 19 cultivars representing 4 species of Capsicum to determine the expression of dormancy in seed germination and emergence.Differences in dormancy were found among cultivars. Fruit age was shown to affect dormancy. Warm, dry storage shortened the length of dormancy while still maintaining a high level of emergence.