Journal of

Seed Science
ISSN 2317-1545
ARTICLE www.abrates.org.br/revista

Stored diaspores of Astronium urundeuva Fr. (M. Allemão) Journal of Seed Science, v.42,
e202042026, 2020
Engl. (Anacardiaceae) submitted to hydropriming
http://dx.doi.org/10.1590/
2317-1545v42236762
Rafael Mateus Alves1* , Monalisa Alves Diniz da Silva2 , Elania Freire da
Silva3 , Robson José Rodrigues Alves2 , Débora Purcina de Moura2 ,
Joyce Naiara da Silva4

ABSTRACT: Seed deterioration is an irreversible process. However, techniques such as

priming have been used after storage, in order to mitigate the harmful effects of aging. This
study aimed to evaluate the physiological performance of A. urundeuva diaspores when
stored and, subsequently, subjected to priming and drying, thus testing the hypothesis that
hydropriming would mitigate deterioration. A completely randomized design was adopted,
in a triple factorial scheme 2x4x4, considering two environments (laboratory and refrigerated
chamber), four storage times (0, 45, 90, and 180 days), and four hydration times (0 h/dry
diaspores, 5 h, 14 h 30 min, and 23 h 30 min). Emergence, speed index, and the number of
days to 50% emergence were assessed, as well as length and dry mass of both shoot and
root systems. The storage of A. urundeuva diaspores for 180 days leads to a reduction in
their physiological performance, regardless of the environment. The hydropriming times are
not effective in attenuating the harmful effects of deterioration. Nevertheless, hydropriming
for 23 h 30 min provides a faster seedling establishment.
*Corresponding author
Index terms: caatinga, aroeira of the hinterland, conservation, controlled hydration. E-mail: rafaelmateusalves@usp.br

Received: 4/17/2020.
Accepted: 6/14/2020.
Diásporos armazenados de Astronium urundeuva Fr. (M. Allemão) Engl.
(Anacardiaceae) submetidos a hidrocondicionamento
Departamento de Produção
1

Vegetal, Escola Superior de

RESUMO: O processo de deterioração das sementes é tido como irreversível, mas técnicas
Agricultura Luiz de Queiroz,
como o priming são realizadas após o armazenamento visando atenuar os efeitos deletérios Universidade de São Paulo (ESALQ-
do envelhecimento. Objetivou-se avaliar o desempenho fisiológico de diásporos de A. USP), 13418-900, Piracicaba – São
urundeuva quando armazenados e, posteriormente, submetidos ao priming e à secagem, Paulo, Brasil.
testando-se, assim, a hipótese de que o hidrocondicionamento funcionaria como um 2
Curso de Agronomia, Universidade
atenuador da deterioração. Adotou-se o delineamento inteiramente casualizado em Federal Rural de Pernambuco -
esquema fatorial triplo 2x4x4, sendo dois ambientes (laboratório e câmara refrigerada), Unidade Acadêmica de Serra Talhada
quatro tempos de armazenamento (0, 45, 90 e 180 dias) e quatro períodos de hidratação (UAST-UFRPE), 56909-535, Serra
(0 h/diásporos secos, 5 h, 14 h 30 min e 23 h 30 min). Avaliou-se emergência, índice de Talhada – Pernambuco, Brasil.
velocidade e número de dias para a ocorrência de 50% de emergência, assim como o 3
Programa de Pós-graduação em
comprimento e a massa seca da parte aérea e do sistema radicular. O armazenamento de Produção Vegetal, (UAST-UFRPE),
diásporos de A. urundeuva por 180 dias acarreta a redução do desempenho fisiológico, 56909-535, Serra Talhada –
independente do ambiente. Os períodos de hidrocondicionamento não são eficientes para Pernambuco, Brasil.
atenuar os efeitos deletérios da deterioração. O hidrocondicionamento por 23 h e 30 min 4
Departamento de Agronomia,
proporciona um estabelecimento mais rápido das plântulas. Universidade Federal da Paraíba
- Centro de Ciências Agrárias (UFPB-
Termos para indexação: caatinga, aroeira do sertão, conservação, hidratação controlada. CCA), 58397-000, Areia – Paraíba,
Brasil.

Journal of Seed Science, v.42, e202042026, 2020

2 R. M. Alves et al.

INTRODUCTION

Astronium urundeuva (Allemão) Engl., Anacardiaceae, is native to northeastern Brazil, where it is popularly known
as aroeira of the hinterland (aroeira-do-sertão). The plant bears pharmacological importance and has the potential for
the restoration of degraded areas, but its survival is at considerable risk due to overexploitation (Centro Nacional de
Conservação da Flora - CNCFlora, 2012).
The production of diaspores in perennial species can vary considerably from year to year (Smith and Samach, 2013),
so that storage is used to regulate their availability (Bewley et al., 2013). The oleaginous diaspores of A. urundeuva do
not display dormancy and have an orthodox storage behavior (Teófilo et al., 2004). They ought to be kept with some
moisture content between 8 and 10% (Guedes et al., 2012). The initial quality might remain the same or decline due
to the deterioration, a process influenced by the moisture content of the diaspores, relative humidity of the air, and
temperature (Marcos-Filho, 2015). Lipid peroxidation is deemed responsible for the membrane damage (Gaschler and
Stockwell, 2017), which is pointed out as the primary characteristic of deterioration (Ratajczak et al., 2019).
In a natural environment, A. urundeuva diaspores quickly lose their viability and vigor after storage (Caldeira and
Perez, 2008). This phenomenon happens because the diaspores are rich in lipids (Guedes et al., 2012), and it tends to
be more intense in surroundings with high humidity (Mira et al., 2015). A widespread consensus state that, although
it can be preserved, seed quality cannot be enhanced during storage (Marcos-Filho, 2015; Mishra et al., 2016). In turn,
priming after storage has been used as an alternative for minimizing detrimental effects resulting from deterioration.
Priming encompasses pre-germinative treatments that benefit seed germination and the subsequent seedling
emergence – this set of techniques is internationally known as “seed enhancements”. Its principle is to supply the seeds
with water under controlled conditions, thus stimulating their metabolism during phases I and II of the imbibition,
without causing the protrusion of the primary root (Bewley et al., 2013). This allows the membrane system to restructure
itself, minimizing the excessive release of exudates (Marcos-Filho, 2015). This method can induce mechanisms of both
protection and repair in seeds, making them capable of tolerating future stresses (Kubala et al., 2015), and favoring
faster germination and the proper establishment of the specimens (Lopes et al., 2019).
Priming can boost germination and improve seedling performance, especially in arid and semi-arid regions,
characterized by irregular rainfall (Finch-Savage et al., 2004). In this context, Meiado et al. (2012) reported that seeds
of species native to the Brazilian Caatinga could only be sown when water is available in the soil (from January to May).
In this case, priming after storage could contribute to extending the sowing period. Similar cases with successful results
include deteriorated seeds of Onobrychis crista-galli (L.) Lam. (Kavandi et al., 2018), which were stored in a germplasm
bank, and were recovered by using osmopriming. In another research, seeds of Trema orientalis Linn. Blume benefited
from a hydration-dehydration process, regardless of whether it were applied before or after storage (Yuniarti et al., 2019).
On that account, this work aimed to assess the physiological performance of diaspores of A. urundeuva when
stored, and subsequently subjected to priming and drying. The hypothesis is that hydropriming works by attenuating
the deterioration.

MATERIAL AND METHODS

Diaspores of A. urundeuva were provided by the Center for Ecology and Environment Monitoring (Núcleo de Ecologia
e Monitoramento Ambiental, NEMA), from the Federal University of Vale do São Francisco (Universidade Federal do
Vale do São Francisco, UNIVASF), in Petrolina, state of Pernambuco (PE), Brazil. They were harvested from the parent
trees in late September and early October 2018, in the city of Salgueiro (PE) (8°03’28” S, 39°05’45” W, 511 m altitude).
Between October and November 2018, the diaspores were manually processed with the aid of a sieve. Next, they were
packed in transparent plastic bags and kept in ambient condition until mid-January 2019, when the experiment initiated.
Storage: the physiological quality of the diaspores was first appraised before they were stored under the specific
experimental conditions (time zero). The diaspores stowed in 250 mL plastic bottles were kept either in laboratory

Journal of Seed Science, v.42, e202042026, 2020

Priming of stored Astronium urundeuva diaspores 3

conditions (32.2±2 °C, 45.67% RH) or inside a refrigerated chamber (20.8±2 °C, 56.16% RH), for 0, 45, 90, and 180 days.
At the end of each period, the moisture content was assessed by the oven method at 105±3 °C for 24 hours. The results
were expressed in percentage (Brasil, 2009).
Hydropriming: after storage, the diaspores were subjected to four hydration times, which were established
according to an imbibition curve obtained previously. Therefore, these times considered the absence of hydropriming
(0h/dry diaspores), and the hydration periods corresponding to ½ of phase I (5 h), ¼ of phase II (14 h 30 min), and ¾
of phase II (23 h 30 min) of the three-phase imbibition process (Lima et al., 2018). For hydropriming, the diaspores
were weighed and then laid onto two blotting-paper sheets, moistened with 15 mL of distilled water (predetermined
value). Then, they were placed inside acrylic gerboxes, at a constant temperature of 25±2 °C, in the presence of
light. After each hydropriming period, two replications with 25 diaspores each had their moisture content analyzed
(Brasil, 2009). At the same time, the remaining diaspores were oven-dried (30±2 °C), and had their weight monitored
every 30 minutes, until reaching approximately a 15% moisture content (Ataíde et al., 2016). Ultimately, seedling
emergence and performance were also evaluated.
Emergence: four replications of 25 diaspores from each treatment were sown in 200-cell plastic trays (18 cm3 per
cell), filled with sterilized sand. The evaluation was carried out daily by counting the normal seedlings (germination
criterion). The tests of emergence speed index (Maguire, 1962) and the number of days to 50% emergence (T50)
(Farooq et al., 2005) were carried out simultaneously.
Seedling length: once the emergence had stabilized ten days past sowing, the seedlings were measured for shoot
length (from the neck to apical meristem) and root system length (from the neck to the extremity of the primary root).
A graduated ruler was used and the results were expressed in cm.seedling-1.
Dry mass of seedlings: the shoot and root system of the normal seedlings of each replication were packed in properly
identified Kraft paper bags. They were dried in a forced-circulation oven set at 80 °C for 24 hours. After that, they were
weighed on an analytical balance with precision to three decimal places, and the results were expressed in g.seedling-1.
Experimental design: the study complied with a completely randomized design, in a triple factorial scheme (2x4x4),
with two environments (laboratory and refrigerated chamber), four storage times (0, 45, 90, and 180 days), and four
hydration times after storage (0 h/dry diaspores, 5 h, 14 h 30 min, and 23 h 30 min).
The analysis of variance employed the F test (p < 0.05), and, once verified a significant interaction, the quantitative
factors were subjected to regression analysis. That not being the case, the averages of the qualitative factors were
compared by the Tukey test (p ≤ 0.05). All statistical analyzes were handled by the software SISVAR v.5.6 (Ferreira, 2011),
and the graphs were built with the software Sigma Plot 10.0.

RESULTS AND DISCUSSION

Neither the emergence nor the performance of A. urundeuva seedlings was affected by the environment, storage
time, or hydropriming. Thus, there was no significant triple interaction, regardless of the variable (Table 1). On the contrary,
the double interaction between the environment and storage period was significant – except for the number of days to
50% emergence (T50). When considered individually, the environment, storage time, and hydropriming factors differed
significantly from T50. Lastly, hydropriming delivered a significant difference as for the lengths of the shoot and root system.
The diaspores had an average initial moisture content of 10.2%. After being stored for 180 days, either under
uncontrolled laboratory conditions or inside the refrigerated chamber, they achieved 11.2% and 11.9%, respectively
(Table 2). Similar results were found by Guedes et al. (2012), who observed that the moisture content of A. urundeuva
diaspores increased throughout storage in all environments tested – but the best conservation result was attained
between 8 and 10%.
The more the diaspore was exposed to the moistened substrate, the higher their average moisture content,
regardless of the storage conditions (Table 2). After the hydroprimed diaspores were dried, their average moisture
content was about 15% (Table 2). According to Ataíde et al. (2016), this moisture level favored the germinative capacity

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4 R. M. Alves et al.

of seeds of Dalbergia nigra (Vell.) Fr. All. Ex Benth (Fabaceae). On the other hand, higher or lower values proved to be
harmful to that species germination.
Regarding the interaction between the environment and time of storage on the emergence (Figure 1) and
emergence speed index (Figure 2), a decrease trend was noticed over time, with the highest values of these attributes
being observed prior to storage (Figures 1 and 2). Even so, the diaspores kept in the refrigerated chamber provided the
highest emergence speed indices (Figure 2). Guedes et al. (2012) considered that the decline in the emergence speed
was the first vigor alteration noticeable in diaspores of A. urundeuva stored for 240 days.

Table 1. Summary of the analysis of variance of the means of the initial development parameters of Astronium
urundeuva seedlings grown from diaspores subjected to the storage, hydropriming, and drying.

Mean Square
Source of variation
DF E ESI T50 LS LRS DMS DMRS
ST 3 13194.7** 36.3** 8.65** 2.59** 1.79** 0.0002 ns
0.001**
SE 1 887.2* 2.85* 3.38* 1.36** 0.02 ns
0.003** 0.003**
HP 3 459.0ns 1.23ns 2.94* 0.35* 0.91** 0.0001ns 0.00008ns
ST x SE x HP 9 48.6ns 0.14ns 0.66ns 0.15ns 0.12ns 0.00007ns 0.0001ns
ST x HP 3 501.7* 1.83* 1.22ns 0.55** 0.7** 0.005** 0.002**
ST x HP 9 152.9 ns
0.41 ns
0.88 ns
0.08 ns
0.04 ns
0.0001 ns
0.00008ns
SE x HP 3 3.69ns 0.06ns 0.55ns 0.08ns 0.12ns 0.0001ns 0.00009ns
Total 127 - - - - - - -
CV (%) 30.9 31.8 18.26 12.5 15.1 29.4 28.4
ns
: non-significant; , : significant at 1% and 5% probability levels, respectively, according to the F test; E - emergence (%); ESI –
** *

emergence speed index; T50 – number of days to 50% emergence; LS – length of the shoot (cm.seedling-1); LRS – length of the root
system (cm.seedling-1); DMS – dry mass of the shoot (g.seedling-1); DMRS – dry mass of the root system (g.seedling-1); ST – storage
time; SE - storage environment; HP - hydropriming.

Table 2. Moisture content of Astronium urundeuva diaspores before and after hydropriming performed subsequent to
storage and drying.

Moisture content (%)

Storage time Hydropriming
Storage environment After drying
(days) Before After
T1 T2 T3 T4 T2’ T3’ T4’
0 10.2 33.9 40.6 54.6 13.8 14.7 14.5
45 11.8 37.8 52.7 56.6 15.6 15.5 16.54
A
90 11.5 34.4 53.7 58.3 13.8 14.9 17.24
180 11.6 36.6 48.3 51.2 15.82 14.6 18.18
0 10.2 33.9 40.6 54.6 13.8 14.7 14.5
45 12.4 40.6 54.1 57.8 15.3 16.5 16.5
B
90 12.1 43.4 56.5 59.7 14.5 16.4 15.7
180 12.8 44.7 50.6 56.9 15.6 15.9 16.6
A: environment without temperature or relative humidity control; B: refrigerated chamber. Seeds without hydropriming (T1); seeds
with hydropriming based on ½ of phase I (T2 – 5 h), ¼ of phase II (T3 – 14 h 30 min), and ¾ of phase II (T4 – 23 h 30 min) of the
three-phase imbibition process.

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Priming of stored Astronium urundeuva diaspores 5

Figure 1. Effect of the times and environments of storage on the emergence of Astronium urundeuva normal seedlings.

Figure 2. Effect of the times and environments of storage on the emergence speed index of Astronium urundeuva
normal seedlings.

The physiological quality of the diaspores stored under uncontrolled environmental conditions was lower, which
corroborates with the findings of Caldeira and Perez (2008). This possibly happens because the average temperature over
the 180 days of storage (32.2±2 °C) accelerated the metabolism and, consequently, intensified the respiration process.
Vigor loss in A. urundeuva diaspores is likely the result of their chemical composition. Despite having an orthodox
behavior, their high oil content causes their germination power to reduce (Teófilo et al., 2004; Guedes et al., 2012),
as lipids have less chemical stability than starches (Balešević-Tubić et al., 2010). Thus, in circumstances where the air
temperature is low, as in a refrigerated chamber (20.8±2 °C), the diaspore metabolism tends to decelerate, resulting in
the decrease of physiological activities and fewer losses due to respiration and lipid peroxidation (Marcos-Filho, 2015).
The mobilization of lipid reserves is paramount during the post-germination until the seedlings properly establish
and become autotrophic. However, the lipid oxidation compromises this process (Hooks et al., 2010). Storage at
low temperatures proved to be an efficient technique to prevent the mobilization of seed reserve compounds in
Amburana cearenses (Allemão) AC Sm. (Fabaceae) (Araujo et al., 2017). It also kept the viability of A. urundeuva
diaspores for up to four months (Oliveira et al., 2018) and guaranteed good seedling emergence of Adenanthera
pavonina L. (Fabaceae) (Gugé et al., 2019).
The diaspores subjected to hydropriming for 23 h 30 min took fewer days to reach a 50% emergence (T50) than those
Journal of Seed Science, v.42, e202042026, 2020
6 R. M. Alves et al.

of the control treatment (dry diaspores) (Figure 3). Consequently, priming provided a positive acceleration in metabolism,
which might represent an ecological advantage in restoration programs for the Brazilian Caatinga. In similar studies,
hydration also favored the emergence of Jatropha curcas L. (Euphorbiaceae) (Braga et al., 2012) and D. nigra. In those cases,
after being subjected to priming, the seeds became physiologically closer to the early phase III of imbibition, which led to
the stimulation of the primary root protrusion (Ataíde et al., 2016). In turn, discontinuous hydration significantly influenced
the number of days to 50% germination of seeds of Mimosa tenuiflora (Willd.) Poir. (Fabaceae) (Lima and Meiado, 2018).
With regards to the storage times, there was an increasing tendency in the number of days to 50% of emergence,
with the storage for 180 days demanding the longest (Figure 4A). As for the storage environments, the diaspores kept
inside the refrigerated chamber required fewer days to deliver a 50% emergence than those stored in the laboratory
environment (Figure 4B).
The length (Figures 5A and 5B) and dry mass (Figures 6A and 6B) of the shoot and root systems showed a statistical

Means followed by the same letter do not differ, according to Tukey’s test (p ≤ 0.05). Seeds without hydropriming (T1); seeds
with hydropriming based on ½ of phase I (T2 – 5 h), ¼ of phase II (T3 – 14 h 30 min), and ¾ of phase II (T4 – 23 h 30 min) of
the three-phase imbibition process.

Figure 3. Effect of hydropriming on the number of days to 50% emergence of Astronium urundeuva normal seedlings.

Means followed by the same letter do not differ, according to Tukey’s test (p ≤ 0.05).
Figure 4. Effect of the times (A) and environments (B) of storage on the number of days to 50% emergence of Astronium
urundeuva normal seedlings.
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Priming of stored Astronium urundeuva diaspores 7

difference for the interaction between environments and storage times. Diaspores stored in the refrigerated chamber
produced more vigorous seedlings, possibly due to the lower intensity of the respiratory process, which delayed the
consumption of reserve substances. Over the storage periods, there was a decline in the variables mentioned above,
especially by 180 days (Figures 5A, 5B, 6A, and 6B). The interference in the translocation of reserves from either the
endosperm or cotyledons to the embryonic axis growth (Amaro et al., 2015) reflects in a reduction of the length and
dry mass of both shoot and root system.
Regardless of the duration, hydropriming did not elongate the shoot or root system, in contrast to the lack thereof. In
turn, priming for 23 h 30 min (T4) favored the growth of the shoot, in contrast to the procedure conducted for 5 h (T2),
and also of the root system, in comparison to the periods of 5 h and 14 h 30 min of hydration (Figure 7). The discontinuous
hydration incremented the development of the A. urundeuva seedlings (Hora and Meiado, 2016). Cycles of hydration
and dehydration in seeds of Antirrhinum spp., Dahlia spp., Impatiens walleriana spp., Salvia splendens Sellow ex Roem. &
Schult. (Lamiaceae), and Zinnia spp. provided seedlings with high amounts of biomass (Ozden et al., 2017).
The minimum moisture content to trigger the germination process depends on the chemical composition and
permeability of the integument (Bradford, 1986). The diaspores of A. urundeuva are oleaginous (Teófilo et al., 2004;

Figure 5. Effect of the times and environments of storage on the length of the shoot (A) and root system (B) of Astronium
urundeuva normal seedlings.

Figure 6. Effect of the times and environments of storage on the dry mass of the shoot (A) and root system (B) of
Astronium urundeuva normal seedlings.

Journal of Seed Science, v.42, e202042026, 2020

8 R. M. Alves et al.

Means followed by the same letter do not differ, according to Tukey’s test (p ≤ 0.05). Seeds without hydropriming (T1); seeds
with hydropriming based on ½ of phase I (T2 – 5 h), ¼ of phase II (T3 – 14 h 30 min), and ¾ of phase II (T4 – 23 h 30 min) of the
three-phase imbibition process.

Figure 7. Effect of hydropriming on the length of the shoot (LS) and root system (LRS) of Astronium urundeuva
normal seedlings.

Guedes et al., 2012), with a resinous mesocarp and a membranous integument (Almeida et al., 1998). Therefore, these
characteristics tend to limit a prompter response to priming, especially when compared to starchy diaspores bearing a
non-waxy permeable integument.
Employing hydropriming for mitigating the effect of the deterioration process during storage still requires further
research, as the final results are deeply influenced by aspects such as the genotype, climatic conditions, pre- and post-harvest
procedures, duration of hydration, drying processes, and the number of hydration-drying cycles. Climate changes also require
knowledge relevant to methods that can be applied to stored diaspores to increase the survival of the seedlings in the field.

CONCLUSIONS

Storing A. urundeuva diaspores for 180 days reduces their physiological performance, regardless of the storage
environment.
Hydropriming, with a discontinuous hydration cycle after the storage, regardless of its duration, is not efficient in
mitigating the deleterious effects of the deterioration process on A. urundeuva diaspores.
A. urundeuva diaspores subjected to hydropriming for 23 h 30 min post storage, regardless of the environment and
storage time, deliver a faster seedling establishment.

ACKNOWLEDGEMENTS

The authors express their gratitude to the Center for Ecology and Environmental Monitoring (Núcleo de Ecologia
e Monitoramento Ambiental, NEMA/UNIVASF), the Project for the Integration of the São Francisco River with the
Hydrographic Basins of the Northern Northeast (PISF), and the Ministry of Regional Development (MDR), for providing
the diaspores of A. urundeuva. Also, to the Federal Rural University of Pernambuco (Universidade Federal Rural de
Pernambuco, UFRPE), Serra Talhada Academic Unit (Unidade Acadêmica de Serra Talhada, UAST), and the Postgraduate
Program in Plant Production (Programa de Pós-Graduação em Produção Vegetal, UFRPE/UAST), for making their
infrastructure available for the execution of this work. To the National Council for Scientific and Technological
Development (CNPq) for granting an academic scholarship to the first author.

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Priming of stored Astronium urundeuva diaspores 9

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