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Acta Scientiarum
ISSN printed: 1679-9283
ISSN on-line: 1807-863X
Doi: 10.4025/actascibiolsci.v36i4.21401

Effect of seed coat on the seed germination and seedling

development of Calophyllum brasiliense Cambess. (Clusiaceae)
Valquria Aparecida Mendes de Jesus1*, Alessandro Lucca Braccini2, Carlos Alberto Scapim2,
Fbio Lcio Santos3, Giovanna Emanuelle Gonalves Mariucci2 and Andria Kazumi
Departamento de Fitotecnia, Universidade Federal de Viosa, Avenida Peter Henry Rolfs, s/n, 36570-000, Viosa, Minas Gerais, Brazil.
Departamento de Agronomia, Universidade Estadual de Maring, Maring, Paran, Brazil. 3Departamento de Engenharia Agrcola, Universidade
Federal de Viosa, Viosa, Minas Gerais, Brazil. *Author for correspondence. E-mail:

ABSTRACT. This work aimed to study the effect of the Calophyllum brasiliense seed coat on the seed
germination process. To this end, three experiments were conducted in laboratory, greenhouse and
screenhouse. From a total of six treatments, five are related to the seed coat (mechanical scarification;
mechanical scarification followed by 2 hours in water, chemical scarification, hot water immersion and
complete seed coat removal) and one control. Laboratory and greenhouse experiments were conducted in a
completely randomized design (CRD). Screenhouse experiment was conducted in a completely
randomized block design (RBD). We evaluated the total percentage, the speed index and the average time
of germination or emergence. Data were subjected to analysis of variance and means compared by LSD
test, at 5%. Under the conditions of this work, it was possible to infer that, in laboratory, mechanical
scarification followed by 2 hours in water increases the proportion and germination speed index (GSI), in
the greenhouse, the complete seed coat removal increases the percentage and emergence speed index (ESI),
and in the screenhouse, mechanical scarification followed by 2 hours in water and chemical scarification
presented the best results. The average germination time was not significantly different in the three
experiments evaluated.
Keywords: native species, diaspore, seed dormancy, Calophyllum brasiliense.

Efeito do envoltrio na germinao da semente e no desenvolvimento de plntulas de

Calophyllum brasiliense Cambess. (Clusiaceae)
RESUMO. Objetivou-se com este trabalho estudar o efeito do envoltrio da semente de Calophyllum brasiliense
no processo de germinao. Foram montados trs experimentos, sendo estes conduzidos em laboratrio, casa de
vegetao e telado. Avaliaram-se seis tratamentos, sendo cinco relacionados ao envoltrio (escarificao mecnica;
escarificao mecnica seguido por 2h em gua; escarificao qumica; imerso em gua quente; e retirada total do
envoltrio) e a testemunha. Os experimentos em laboratrio e casa de vegetao foram conduzidos em
delineamento inteiramente casualizado. No telado, o experimento foi conduzido em delineamento em blocos
casualizados. Avaliaram-se a percentagem total; o ndice de velocidade e o tempo mdio de germinao ou
emergncia. Os dados foram submetidos anlise de varincia e as mdias comparadas pelo teste t (LSD) a 5% de
probabilidade. Foi possvel inferir que no laboratrio, a escarificao mecnica, seguido por 2h em gua, aumenta
a percentagem e o ndice de velocidade de germinao (IVG); na casa de vegetao, a retirada total do envoltrio
aumenta a percentagem e o ndice de velocidade de emergncia (IVE); e no telado, a escarificao mecnica,
seguida por 2h em gua, e a escarificao qumica apresentam os melhores resultados. O tempo de germinao
no apresentou diferena significativa nos trs experimentos avaliados.
Palavras-chave: espcie nativa, disporo, dormncia, Calophyllum brasiliense.

Introduction Calophyllum brasiliense Cambess., commonly

known as guanandi, a climax species (SANTOS
Family Clusiaceae includes trees, bushes,
et al., 2008), is a large arboreal plant, with height
lianas and herbs of economic interest for timber ranging from 20 to 30 m, diameter between 40 and
production, edible fruit, chemical derivatives of 60 cm and wood density of 0.62 g cm-3 (LORENZI,
pharmaceutical interest and others. Among the 2002). In Brazil, it occurs naturally from the
genera, there is Calophyllum L., which produces Amazon region to the north of Santa Catarina State,
hardwoods (JNIOR et al., 2005). and is mainly found in the Atlantic rainforest
Acta Scientiarum. Biological Sciences Maring, v. 36, n. 4, p. 451-456, Oct.-Dec., 2014
452 Jesus et al.

(CARVALHO, 1994; LORENZI, 2002; NERY each experiment, they were subjected to six
et al., 2007a; REIS et al., 2009). treatments, five related to coat and one to the
In addition to presenting chemical and untreated control.
pharmacological importance, as reported by Jnior
et al. (2005) and Noldin et al. (2006), it is Treatments
recommended for riparian forest restoration, mostly Control treatment: seeds with intact coat,
in places subjected to periodic flooding from without treatment after processing.
medium to long term (CARVALHO, 1994). This Mechanical scarification: the treatments using
species has good quality wood and characteristics mechanical scarification submitted the seeds to a
similar to the mahogany and cedar, providing a crack in their coat. Thus, a hammer was used to
possible alternative to replace them. break the seed coat without any damage to the
The first vegetative stage of the plant after seed embryo.
germination is known as seedling, which has Chemical scarification for 15 min.: seeds were
enormous value for the study of population immersed in concentrated sulfuric acid for 15 min.
dynamics, forestry, seed storage, works in nurseries To perform this procedure, we used a beaker and a
and in forest conservation and restoration (SOUZA, glass rod, under a lab fume hood.
2003). According to Masetto et al. (2008), Immersion in hot water: consisted of immersing
information regarding the positive characteristics seeds in hot water at boiling point for one min.
about seed germination, through which it is possible Then, the seeds were immersed in water at ambient
to achieve a higher germination rate, although temperature for 48 hours.
essential, is scarce in the literature. Procedures to Mechanical scarification and water for 2 hours:
conduct germination test for most native species are this treatment consisted of the mechanical
still limited in the Rules for Seed Analysis (BRASIL, scarification followed by immersing the seed in
2009). water at ambient temperature for two hours.
Therefore, given the importance and potential of Total removal of seedcoat: the treatment
Calophyllum brasiliense (Clusiaceae) and the lack of consisted of completely removing the seed coat,
information, standardization, and analysis methods adopting a procedure similar to mechanical
for the vast majority of forest species, especially for scarification. However, after cracking, the whole
the species in question, this study evaluated the seed coat was removed manually.
effect of Calophyllum brasiliense seed coat on the seed
Experiment 1
germination process, in order to achieve a greater
percentage of germinated seeds in a faster and The germination test was conducted in the
uniform process. absence of light in a germination chamber (Tecnal,
TE-404), under constant temperature of 30C. This
Material and methods temperature is considered optimal for C. brasiliense
germination (FERREIRA et al., 2007; NERY
Seeds of Calophyllum brasiliense Cambess. et al., 2007b).
(Clusiaceae) were obtained from artificial Paper towels were used as substrate, in a roll
populations containing trees between four and six form, moistened with distilled water (2.5 times the
years old, in Monte Alto City, So Paulo State. They dry paper mass), according to the Rules for Seed
were similar, originated from a population Analysis (BRASIL, 2009). Observations were made
containing only trees of C. brasiliense. Seeds daily, during 90 days. Seeds were considered
presented a 25% initial average humidity, which was germinated when showed radicle protrusion and/or
determined by the oven method at 105 3C for shoot equal to or exceeding 2 mm. The laboratory
24 hours, using a mechanical oven with forced air experiment was performed using a completely
circulation, based on the water mass of seeds. randomized design, which consisted of four
After selected, seeds were aseptically treated with replications with 25 seeds per each treatment.
bleach commercial solution containing sodium
hypochlorite at 2.0%. Seeds were immersed in this Experiment 2
solution for two min. and rinsed in distilled water At the greenhouse, not climatized, germination
five times. test was conducted on plastic trays with coarse sand
To evaluate the seeds germination, three as substrate. During the germination test, minimum
experiments were set up, which were developed one temperature of 7C and maximum of 44C was
in the laboratory, one in the greenhouse and one in observed. Treatments were watered twice a day, in
the screenhouse. To study the effect of seed coat in the early morning and late afternoon. Observations
Acta Scientiarum. Biological Sciences Maring, v. 36, n. 4, p. 451-456, Oct.-Dec., 2014
Effect of the Calophyllum brasiliense seed coat on the seed germination process 453

were made daily, during 90 days. Seeds were There was no significant difference (p > 0.05)
considered germinated when produced seedlings between control and treatment with immersion in
with all essential structures to the proper plant hot water for 48 hours (Table 1), which presented
development in the nursery. To characterize the the lowest germination percentage. This result is
seedling, we used the concept proposed by Souza due to the immersion in hot water, which probably
(2003), which establishes that a well-developed caused a seed coat rupture or damage to the embryo.
seedling must have the following structures: primary Nery et al. (2007a) studying C. brasiliense seeds,
root, stem base, hypocotyl, epicotyl and eophylls. found the necessity of removing the seed coat to
The greenhouse experiment was installed using a occur imbibition, an essential fact for the
completely randomized design, consisted of four germination process.
replications with 25 seeds per each treatment.
Table 1. Total average percentage of germination and
Experiment 3 germination speed index of Calophyllum brasiliense (Clusiaceae)
seeds subjected to different treatments in laboratory.
The screenhouse is a structure fully covered with
transparent plastic, and free sides, which softened Treatments Germination (%) GSI
Control 5E 0.04 D
the temperature. Germination test was conducted Mechanical scarification 62 C 0.45 C
using seed beds, with four areas of 1 m2 each, Chemical scarification - 15 min. 77 B 0.58 B
Immersion in hot water - 48 hour 5E 0.03 D
corresponding to the experiment blocks. Seeds were Mechanical scarification and water for 2 hours 92 A 0.72 A
placed on the local soil and covered with manure, in Total seed coat removal 44 D 0.39 C
order to maintain moisture. Each block was watered Means followed by the same letter, in the column, do not differ by LSD test, at 5%
probability. GSI: germination speed index.
twice a day, in the early morning and late afternoon.
Observations were made daily, also during 90 days. Seed coat rupture or even its complete removal
Seeds were considered germinated when seedlings promoted a significant increase in seed germination
presented all essential structures to the proper plant (Table 1). According to Marcos Filho (2005), seed
development. Screenhouse experiment was coat regulates the seed imbibition speed, controls gas
conducted in a randomized block design, consisted exchange, causes dormancy and may also prevent
of four replications and 25 seeds per treatment in seed deterioration, acting as a barrier to
each block. microorganisms entry. This result differs from the
Evaluation and data analysis results found by Nery et al. (2007b), in which the
final germination percentage was not different
The treatments were evaluated through seed between C. brasiliense seeds with and without seed
germination (G) or seed emergence (E),
germination speed index (GSI) or emergence speed
Among the following treatments: mechanical
index (ESI) and average germination time (AGT) or
scarification, chemical scarification treatment,
average emergence time (AET) of C. brasiliense seeds.
mechanical scarification followed by immersion in
To calculate the GSI and average germination time,
water for 2 hours and total seed coat removal, there
we used the data obtained in germination tests. GSI
was no significant difference (p > 0.05). In
was calculated using the equation of Maguire
laboratory, the high humidity inside the germinator
(1962), suggested by Nakagawa (1999). The average
germination time was calculated according to chamber besides the higher water amount favored
Santana and Ranal (2004). the proliferation of fungi and microorganisms,
Data for total germination, GSI or ESI, and directly affecting seeds without coat due to
average germination time or average emergence susceptibility (Table 1). Sulfuric acid acts on the
time, were subjected to analysis of variance. Means seed coats removing the outer layers, such as cuticle
between treatments were compared by LSD test at and the cuticle layers, throughout the coat, allowing
5% probability. All analyses were run using the more homogeneous permeability degrees
statistical program Sisvar (FERREIRA, 2006). (SANTARM; QUILA, 1995). In contrast, in
mechanical scarification, seed coat is broken only in
Results and discussion a small area, so it does not damage the embryo. In
this way, the seeds mechanically scarified present
Experiment 1 only a small area allowing easy water entry, while
The analyses of variance performed for chemically scarification may have an advantage in
germination and germination speed index in water absorption by presenting a more uniform
C. brasiliense seeds indicated significant differences at scraping along the entire coat, facilitating and
5% among the treatments. uniformizing water absorption. This may be the
Acta Scientiarum. Biological Sciences Maring, v. 36, n. 4, p. 451-456, Oct.-Dec., 2014
454 Jesus et al.

reason for the higher germination percentage in Experiment 2

chemically scarified seeds compared with those In the greenhouse, no significant difference was
mechanically scarified (Table 1). On the other hand, detected between the control and hot water
seeds scarified mechanically and placed in immersion treatment (Table 2). The no rupture of
immediate contact with water for 2 hours showed a the seed coat probably acted hindering the
higher germination percentage. C. brasiliense seeds absorption of water by the seed. Water absorption is
are recalcitrant, as evidenced by Carvalho essential for the germination; therefore the seed coat
et al. (2006); so, seeds are sensitive to desiccation, effect of such treatments resulted in the lowest
demonstrating higher percentages of germination in accumulated percentage of emergence of
immediate contact with water, after scarification, C. brasiliense seedlings. Hot water immersion
probably because such a procedure has maintained a treatment may also have caused damage to the
high water content in the seeds and has contributed embryo, affecting seedling emergence.
to a more uniform absorption. Water contact after
scarification is an alternative to C. brasiliense, since Table 2. Total average percentage of emergence and emergence
speed index of Calophyllum brasiliense (Clusiaceae) seeds subjected
after scarification, the permanence of seeds in water to different treatments in greenhouse.
probably kept a high water content in seeds and
Treatments Emergence (%) ESI
caused a uniform absorption, and resulted in a Control 6D 0.04 D
higher germination percentage (Table 1). Mechanical scarification 33 C 0.24 C
Chemical scarification - 15 min. 69 B 0.45 B
On average, the results obtained for the GSI Immersion in hot water - 48 hour 8D 0.06 D
(Table 1) showed no significant difference between Mechanical scarification and water for 2 hours 65 B 0.45 B
Total seed coat removal 83 A 0.58 A
intact seeds (control) and seeds that received the
Means followed by the same letter, in the column, do not differ by LSD test, at 5%
immersion in hot water. The difficulty in absorbing probability. ESI: emergence speed index.
water, due to the seed coat, was probably responsible
for the lower values of GSI. Mechanical scarification However, in the other treatments, there was a
and total seed coat removal were not significantly significant increase in the emergence percentage.
different to each other (p > 0.05). The low GSI The increase in the emergence percentage can be
found in seeds subjected to mechanical scarification explained by the seed coat rupture, facilitating the
is due to an insufficient water absorption caused by a water absorption by the seed, resulting in an
small rupture, thus affecting the germination rate. increased germination percentage. The results
On the other hand, the complete removal of the indicated that the permanence of the mechanically
seed coat may have caused an excessively rapid water scarified seeds in water significantly improved
imbibition. Seeds chemically treated were emergence percentage. Chemical and mechanical
significantly different (p < 0.05) from other scarifications followed by 2 hours in water
treatments. Chemical scarification allowed water immersion were not significantly different
uptake throughout the seed coat, which positively (Table 2). The lack of difference between these
influence the germination rate. The best treatments is likely to be related to the efficiency of
germination speed index results in laboratory for chemical scarification in this study, allowing
C. brasiliense seeds was found in the mechanical adequate water uptake by the seed, so that these
scarification treatment, followed by seeds remaining results did not differ to each other. Nevertheless, the
in water for 2 hours, as presented in Table 1. no rupture of the seed coat limited water uptake,
There were no significant differences (p > 0.05) thus limiting the emergence percentage.
between treatments for average germination time. The complete removal of seed coat resulted in a
Seed coat rupture, although promoting a higher higher emergence percentage, due to the easier
germination percentage and a higher germination water absorption by the seed, evenly, facilitating the
speed index, showed no significant difference in germination process. In the greenhouse experiment,
average germination time between treatments with it was observed that the water percolates through the
or without seed coat rupture. According to Ferreira sand, thus preventing its accumulation that harms
and Borghetti (2004), the average time of the germination and favor microorganism attack.
germination is often directly related to the species, Additionally, unlike the laboratory experiment, with
characterizing the strategies of each species to humidity approximately to 100%, favoring attack of
become established in the environment as soon as seeds by fungi and other microorganisms, the
possible. Therefore, seeds of C. brasiliense showed no greenhouse with sand substrate and total seed coat
significant difference when comparing the different removal favorably affected the germination
treatments related to seed coat. percentage. The sand facilitates water percolation,
Acta Scientiarum. Biological Sciences Maring, v. 36, n. 4, p. 451-456, Oct.-Dec., 2014
Effect of the Calophyllum brasiliense seed coat on the seed germination process 455

favoring the water absorbtion by the seed and it is substrate. Probably, this retention resulted in a rapid
not conducive to fungi development and imbibition, affecting the germination process and
microorganisms, as occurs in environments with the high water content favors the proliferation of
high humidity. fungi and microorganisms, which mainly affect the
The analysis of variance with data of emergence seeds without coat because of their susceptibility.
speed index determined in the greenhouse
evidenced a significant difference (p < 0.05) among Table 3. Total average percentage of emergence and emergence
speed index of Calophyllum brasiliense (Clusiaceae) seeds subjected
the treatments. Thus, the means for the emergence to different treatments in screenhouse.
speed index were compared by LSD test at 5%
Treatments Emergence (%) ESI
probability and the results are presented in Table 2. Control 4D 0.03 D
There was a similar behavior among treatments Mechanical scarification 63 B 0.42 B
for the emergence percentage and emergence speed Chemical scarification - 15 min. 74 A 0.51 A
Immersion in hot water - 48 hour 7D 0.05 D
index in the greenhouse. In Table 2, treatments Mechanical scarification and water for 2 hours 78 A 0.50 A
related to the seed coat rupture promoted a higher Total seed coat removal 31 C 0.23 C
emergence speed index. Similar results to those Means followed by the same letter, in the column, do not differ by LSD test, at 5%
probability. ESI: emergence speed index.
obtained in this work were found by Nery
et al. (2007b), who reported for C. brasiliense, higher Mechanical scarification showed significantly
GSI values in seeds without coat, and noted a better results when compared with seeds without
possible coat interference with water uptake by the seed coat. This result is due to the function of seed
seed, which slows germination and causes coat as a protection to the embryo. According to
dormancy. The analysis of variance performed for Santos et al. (2009), the coat acts as seed protection,
the average emergence of C. brasiliense seedlings in so that its absence becomes a limiting factor.
the greenhouse showed no significant differences at According to the results in Table 3, there was no
5% probability among the treatments. The difference between seeds that were mechanically
treatments applied to the seed coat of C. brasiliense scarified followed by water immersion for 2 hours
did not affect the average emergence time probably and chemically scarified seeds. Both treatments
because it is a survival mechanism of the species. resulted in greater seedling emergence in the
screenhouse, which is due to enough water
Experiment 3 absorption, due to seed coat rupture, thus favoring
The results obtained in this experiment showed the germination process, compared with other
the species development subjected to conditions treatments. At the same time, the seed coat
very close to the field, however with partial permanence worked as protection for the embryo,
protection by the effect of the clear plastic cover, water output control by the seed and possibly against
which softens the direct effects of weather. attack by fungi and/or predation.
The analysis of variance for treatments The analysis of variance for the emergence speed
performed in the screenhouse indicated index determined in screenhouse is presented in
non-significant differences between blocks used in Table 3. No significant differences were observed
the experiment. This result is probably due to the between blocks used in the experiment. This fact is
uniformity of the area where the experiment was due to the uniformity of the area where the
conducted. However, we observed significant experiment was set up. However, there were
differences between treatments, and the means for significant differences between treatments analyzed,
percentage emergence were compared by LSD test, and the means for the emergence speed index were
at 5% probability. compared by LSD test, at 5% level of probability.
Based on the results presented in Table 3, we The results for the emergence speed index in the
found higher seedling emergence percentages in screenhouse were similar to the results of
seeds subjected to treatments related to the seed coat emergence in the same conditions (Table 3). Similar
rupture. As previously described, rupture favors the results were found by Cardoso and Pereira (2008) in
water uptake by the seed, favoring germination a study on the effect of water potential on seed
process. Meanwhile, seeds with complete seed coat germination of Drymaria cordata. These authors
removal, despite showing significantly higher observed that the treatments that affected the
emergence rates when compared to intact seeds or germinability also affected the germination speed,
seeds with hot water immersion had a lower showing similar behavior. The similarity in behavior
emergence percentage in the screenhouse. This low between the total emergence percentage and ESI is
emergence percentage can be explained by the ability probably related to the same limiting fator; in this
of soil water retention when compared with sand study, it is believed that water was the limiting factor.
Acta Scientiarum. Biological Sciences Maring, v. 36, n. 4, p. 451-456, Oct.-Dec., 2014
456 Jesus et al.

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License information: This is an open-access article distributed under the terms of the Creative
evaluation for seedling emergence and vigor. Crop Commons Attribution License, which permits unrestricted use, distribution, and reproduction
Science, v. 2, p. 176-177, 1962. in any medium, provided the original work is properly cited.

Acta Scientiarum. Biological Sciences Maring, v. 36, n. 4, p. 451-456, Oct.-Dec., 2014