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PH and gibberellic acid overcome dormancy of seeds of Brachiaria brizantha

cv. Marandu

Article · January 1999

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Revista Brasileira de FisiologiaVegetal, 11(1):51-54, 1999.

COMMUNICATION

pH AND GIBBERELLIC ACID OVERCOME

DORMANCY OF SEEDS OF Brachiaria brizantha CV.
MARANDU 1

Henrique Duarte Vieira2, Roberto Ferreira da Silva3, Raimundo Santos

Barros4
Laboratório de Fitotecnia-CCTA, Universidade Estadual do Norte Fluminense, Campos
dos Goytacazes, RJ, 28015-620.

ABSTRACT: Freshly harvested seeds of Brachiaria brizantha cv. Marandu were cleaned and dried to
12% of moisture content. The seeds were manually dehulled and treated with sodium hypochlorite 0.5%
for five minutes. They were then infiltrated under vacuum for 10 minutes with solutions of pH 2, 3, 4, 5, 6
and 7 and gibberellic acid (pH 2.0 e 6.0). Germination was monitored at 30oC and seeds were considered
as germinated when, after 120 hours, then showed protrusion of the radicle. The physiological dormancy
of the seeds was partially released by low pH solutions and by gibberellic acid. The combination of low pH
solution with gibberellic acid showed an additive effect in the promotion of the germination of dormant
seeds.
Additional index terms: braquiarão, grass, pasture.

pH E ÁCIDO GIBERÉLICO NA SUPERAÇÃO DA DOMÊNCIA EM SEMENTES DE

Brachiaria brizantha CV. MARANDU

RESUMO: Sementes recém-colhidas de braquiarão (Brachiaria brizantha cv. Marandu) foram bene-
ficiadas e secas até 12% de umidade. As sementes foram descascadas manualmente e tratadas com
hipoclorito de sódio a 0,5%, por cinco minutos. Em seguida, foram infiltradas a vácuo por 10 minutos,
com as soluções de pH 2, 3, 4, 5, 6 e 7 e de ácido giberélico (pH 2,0 e 6,0). Foram consideradas
germinadas as sementes que, após 120 horas em câmara de germinação a 30oC, apresentaram protrusão
da radícula. A dormência fisiológica das sementes foi liberada sob maior acidez. A combinação de pH
baixo com ácido giberélico mostrou um efeito aditivo na promoção da germinação de sementes dormen-
tes.
Termos adicionais para indexação: braquiarão, dormência, gramínea, pastagem.

1 Received 20/02/1999 and accepted 11/05/1999.

2 Eng. Agr. DS, Laboratório de Fitotecnia, CCTA, UENF, Campos

dos Goytacazes, RJ, 28.015-620, e-mail: henrique@uenf.br

3 Eng., Agr., PhD., Prof., Laboratório de Fitotecnia, UENF - Cam-

pos dos Goytacazes, RJ, 28.015-620, e-mail:roberto@uenf.br

4 Eng. Agr. PhD. Prof.,Depto. de Biologia Vegetal, UFV, Viçosa,
MG, 36.570-000, e-mail:rsbarros@mail.ufv.br

51
 52
INTRODUCTION
For pasture establishment with braquiarão seeds their
undesirable low germination has been attributed to a high
degree of dormancy (Garcia & Cícero, 1992). The
germination of seeds of the genus Brachiaria has been
investigated by several authors (Grof, 1968; González et
al., 1994; Voll et al., 1997). Grof (1968) and Renard &
Capelle (1976) observed that seed covers, gluma, palea
and lemma constituted a barrier to the germination, due
to impermeability to water and oxygen. Besides the
dormancy caused by seed covers, an endogenous
dormancy is supposed to block germination of freshly
harvested seeds. This endogenous or physiological
dormancy could be associated with the balance of growth
regulators in the seed (Khan, 1971; Bewley & Black,
1994). In B. brizantha cv. Marandu, Garcia & Cícero (1992)
observed that the treatment of the seeds with sulfuric
acid promoted 15 to 18% germination, while gibberellic
acid caused 14% germination.
The action of low pH of the sorrounding medium in
the germination of dormant seeds was observed in rice
by Roberts (1963), in barley and celery by Palevitch &
Thomas (1976) and in red rice by Footitt & Cohn (1992).
According to Roberts (1963) the effect of the low pH on
germination was due to an increase of free H+ ions,
affecting some physiological phenomena. Bewley & Black
(1994) believe that a higher availability of protons could
result in a larger extensibility coeficient of the cell walls,
leading to the growth of the cells. The pH effect has also
been associated with gibberellins in lettuce seeds, in
which low pH seems to have increased their sensitivity
to the gibberellins, while in celery the low pH seemed to
have stimulated a greater penetration of the gibberellins
into the cells (Hsiao et al., 1984; Palevitch & Thomas,
1976). Footitt et al. (1995) observed that the release of
the dormancy of seeds of red rice was directly associated
to the acidification of the embryo tissues.
McLean & Grof (1968) found that the acid
escarification of seeds of Brachiaria ruziziensis was more
efficient than the mechanical one in the release of the
dormancy, suggesting that low pH could also affect the
physiological dormancy of seeds in the genus Brachiaria.
The objective of this work was to evaluate the effects
of gibberellic acid and pH of the surrounding medium
separetely or in combination on the germination of
dormant seeds of Brachiaria brizantha cv. Marandu.
MATERIAL AND METHODS
The experiments were conducted in the Plant Science
Laborator y of Universidade Estadual do Nor te
Fluminense, Campos dos Goytacazes, Rio de Janeiro,
Brazil. Braquiarão seeds were produced in a pasture
located at Mimoso do Sul, Espírito Santo, Brazil.
The braquiarão (Brachiaria brizantha cv. Marandu)
seeds were harvested from plants in a pasture system.
Harvests occurred from February through July 1997,
when no seeds were observed attached to the clusters
of the plants. For harvesting, the seeds were released
R. Bras. Fisiol. Veg., 11(1):51-54, 1999.
Vieira et al.
fallen down from the bunches following a mild impact to
the inflorescence and caugh before falling to the ground.
This constituted an advantage since the floral banners
were not damaged and facilitated subsequent harvests
in the period of seed production.
Seeds were then cleaned with a pneumatic blower;
afterwards the remaining sludges were manually
removed for obtaining pure seeds. Seeds were dried to
11-12% moisture content in a ventilated oven.
Subsequently, seeds were stored in paper bags in the
laboratory until the experiments.
Before the treatments, seeds were individually
dehulled with a forceps to remove glumes, lemma and
palea, leaving only the true caryopses. They were then
treated with sodium hypochlorite 0.5% for five minutes
and thoroughly rinsed with distilled water.
The seeds were immersed in the treatment solutions
in a beaker and submitted to a vacuum for ten minutes,
in the following way: three minutes of vacuum, followed
by three minutes without vacuum, and a further four
minutes of vacuum, for greater penetration of the solutions
into the seeds (Vieira & Barros, 1994). Then, seeds with
the solutions were transferred to 100 mm X 15 mm Petri
dishes, containing a layer of blotting paper. Petri dishes
and the blot paper were sterilized previously at 140oC for
two hours.
The solutions of different pH (MacIlvaine’s buffer, 10
mM) and those containing the gibberellic acid at pH 2.0
and 6.0 were renewed twelve hours from the start of the
experiments, so that the initial pH could be maintained.
Seeds were maintained in a germination chamber at
30oC for 120 hours. Germination was evaluated by the
protrusion of the radicle, according to Mott et al. (1976).
A completely random design was used with four
replications; each replication was composed of twenty
seeds. For statistical analysis, the data were transformed
to Arcsin (%G/100)1/2, so that they could be adjusted to a
normal distribution (Steel & Torrie, 1980). Duncan’s test
at a level of 5% was used to estimate the significance of
differences among means of the treatments.
RESULTS AND DISCUSSION
The effect of the pH on the dormancy of braquiarão
seeds is shown in figure 1. As solution pHs depart from
neutrality towards acidity, an increase is observed in the
germination of the dormant seeds; at pH 2.0, for instance,
germination was about 65%, significantly different of
germination at pH 3.0, which was, in turn, different from
the germination at pH 4.0 and 5.0. The effect of low pH
on germination was observed by Roberts (1963) in seeds
of rice: as pH of the solution decreased, the germination
percentage increased. He suggested that germination
was favored by an increase of H+ concentration. This
increase in H+ concentration could lead to an acidification
of the cell walls, reducing their rigidity, and allowing a
greater cell growth (Bewley & Black, 1994). Adkins et al.
(1985) also found that the dormancy of seeds of wild oat
(Avena fatua) was released by the lowering of the pH.
Among several explanations, low pH could be promoting
 pH and gibberellic acid . . . 53

FIGURE 1 - Effect of pH on
germination of dormant seeds of
Brachiaria brizantha cv. Marandu,
24 days after harvesting. The
bars represent the mean
standard errors. Mean values
followed by the same letter were
not significantly different (P =
0.05) by Duncan’s test.

an increase in the diffusion of inhibitors from the cells,
as suggested by Cohn et al. (1987). In fact, Footitt &
Cohn (1992) found a decrease in the pH of the embryo
cells before germination of dormant seeds of red rice
occurred; no pH change was observed in embryo cells
of seeds which remained dormant.
In figure 2, the effect of gibberellic acid combined with
low pH can be observed. Solutions of pH 2.0 and
gibberellic acid (pH 6.0) were able to release significantly,
at a level of 5%, the physiological dormancy of the
braquiarão seeds. At pH 2.0, gibberellic acid promoted
germination of an additive kind as compared to each factor
separately. It thus seems that each factor was acting upon
dormancy in a different way. Seeds of wild oat also
exhibited a response to both low pH and gibberellic acid
(Adkins et al., 1985). Similar results were observed in
seeds of Apium graveolens, by Palevitch & Thomas
(1976). According to the authors the effect of the pH could
be explained in two ways. The low pH could have facilitated
the penetration of the gibberellins through the membrane

FIGURE 2 - Effect of the pH and

of the gibberellic acid (0.1
mol.m -3 ) on germination of
dormant seeds of Brachiaria
brizantha cv. Marandu, with 22
days after picking. The bars
represent the mean standard
errors. Mean values, followed by
the same letter were not
significantly different (P = 0.05) by
Duncan’s test. Full rectangule
represents the summation of the
effects of pH 2.0 and gibberellic
acid.

R. Bras. Fisiol. Veg., 11(1):51-54, 1999.

 54
lipid constituent since they were in a protonated form.
Alternatively, the increase in hydrogen ion concentration
might have lead to changes in the cell wall, thus allowing
growth, a phenomenon known as acid effect. In dormant
seeds of braquiarão, it is possible that another mechanism
was active since pH 2.0 or gibberellic acid alone was
able to cause dormancy break and, as mentioned, the
simultaneous action of both was additive. This did not
occur in celery seeds, in that low pH by itself did not
promote germination. In this case it is likely, that the
synthesis of endogenous gibberellins was stimulated by
low pH. This factor could also promote the lixiviation of
inhibitors in the embryonic tissues. Some mechanisms
observed by Palevitch & Thomas (1976), in celery seeds,
could also be acting in the dormant seeds of braquiarão.
In conclusion, the physiological dormancy of
braquiarão seeds was partially released by low pH and
by gibberellic acid separately. The gibberellic acid supplied
in solution at pH 2.0 showed an additive effect with this
pH in the increase of germination of dormant seeds of
braquiarão.
ACKNOWLEDGEMENTS
To FAPERJ and UENF.
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