QUALITY OF SEED PRODUCED BY TROPICAL FORAGE LEGUMES ON LOW FERTILITY SOILS | Amar | AGROLAND: The Agricultural Sciences Journal 8120 26654 1 PB
Agroland: The Agriculture Science Journal, 2016 June 3(1) : 1– 13
ISSN : 2407 - 7585
E-ISSN : 2407 - 7593
QUALITY OF SEED PRODUCED BY TROPICAL FORAGE LEGUMES
ON LOW FERTILITY SOILS
Andi L. Amar1), Robert A. Congdon 2), Christopher P. Gardiner3), Ross J. Coventry2)
1)
Faculty of Animal Husbandry and fisheries. University of Tadulako. Palu. 94117 Central Sulawesi. Indonesia.
2)
School of Marine and Tropical Biolog James Cook University. Townsville. Queensland. 4811Australia.
3)
School of Veterinary and Biomedical Science James Cook University. Townsville. Queensland. 4811Australia.
ABSTRACT
Soil fertility can influence seed production and seedling growth. This study compared seed
yields, seed and seedling characteristics of eight forage legumes (Stylosanthesscabra, S. hamata,
Centrosemabrasilianum, C. pascuorum, Desmanthuspubescens, D. Virgatus, Macroptiliumbracteatum
and M. martii) grown on two soil types (red and yellow kandosols), of low fertility, that are
widespread in the Australian tropical rangelands. Seeds of the Desmanthus species germinated most
rapidly and, with M. bracteatum, produced the highest proportion of readily germinable seed. M.
bracteatum seed produced on the Yellow Kandosol germinated significantly faster, as did the S.
hamata from the Red Kandosol than the other species. S. scabra cv. Seca and C. brasilianum
produced more readily germinable seeds from plants grown on the Red Kandosol. C. pascuorum,
D. pubescens and M. bracteatum produced a higher percentage of readily germinable seeds and
fewer hard seeds when grown on the Yellow Kandosol. No significant differences were found in
seedling vigour or biomass allocation from seeds produced on the different soil types.
Key Words : Fertility, forage, legumes, seed, soil, quality.
INTRODUCTION
It is well recognised that plant
development and seed production are
influenced by the surrounding environment.
The environmental conditions during seed
development, along with genetic characteristics
and agronomic techniques have considerable
effect on seed yield and components
of yield through their effect on plant
reproductive physiology (Martiniello, 1998).
The quality of the seed sown, including its
vigor, determines the success of the resulting
plants; while seed quality is reflected in
seedling density and vigor, uniformity of
crop growth, and plants competitiveness, as
well as forage production insome species
(Hampton, 1995). Many publications have
demonstrated that environmental experiences
(e.g. light, nutrient, temperature, water,) of
the maternal plant have influences on
the performaces of the resulting progeny
(Rossiter, 1996; Galloway, 2002; Etterson
and Galloway, 2002; Galloway, 2005;
Luzuriaga, et al., 2006; Galloway and
Etteson, 2007; Uller, 2008; Wolf and Wade,
2009; Mitrovic, et al., 2010).
Nutrient supplies from the soil
in which a plant grows is often found to
be an important factor contributing to
environment-induced maternal effects.
Nutrient status of a plant may affect the
number, size and quality of seeds produced,
as well as the growth and competitive
ability of the plants progeny (Aarssen and
Burton 1990, Sills and Nienhuis 1995).
This is in agreement with Miao et al.
(1991a, 1991b) who showed that the
maternal nutrient environment influences
the competitive ability of progeny in
Plantago spp.; and Galloway (2001)
proved this with a strong effect of maternal
nutrient environment on the germination of
offspringin herbaceous plant Campanula
americana . Though, this is not always
the case. Luzuriaga, et al., (2006) found
1
that added nitrogen to Sinapis arvensis
plants only slightly reduced the germination
rate of its seeds, but not the germination
percentage.
A plant‟s ability to grow under
certain conditions is not enough to indicate
its ability to persist in an ecosystem for
long periods, as is required for a pasture
species. Production of a large number of
seeds is an important factor controlling
plant persistence, as is the seed's capacity
to germinate, to provide new plants for
regeneration. This is particularly important
for an annual species whose persistence
is strongly dependent on both seed production
and viability. Furthermore, establishment
of a good sward is also an important
criterion for pasture production. Early
seedling development plays an important
role in the long-term growth of a plant,
and in the dynamics of plant populations
in the field (Hendrix 1984; Schmitt and
Antonovics 1986). Several studies have
demonstrated the importance of early
emergence in relation to the competitive
ability of the resulting seedlings (Crawley
and Nachapong 1985; Dolan 1984).
Seeds resulting from a previous
study entitled “an Early Growth Study”
(Amar et al. 2012) were tested toinvestigate
the effects of plant genotype and soil type
on the performance of the progeny of the
studied legumes. Although not as strong as
genotype effects, it was found from the
early study that the soil type significantly
affected herbage yield, time to flowering
and the number of seed produced (Amar
et al., 2012). Therefore, it is hypothesised
that edaphic influences on the parent
plants may further affect the viability of
the seeds produced and the performance
of the resulting seedlings. Consequently,
two experiments were conducted here, as a
„Germination Study‟ and a „Seedling Study‟
which are described below.
The „Germination Study‟ compared
seed germination over time, the proportions
of readily germinable seeds, hard seeds,
and non-viable seeds resulting from the 8
legume genotypes grown on the different
soils. The germination criteria, however,
may not be sufficient to judge seed quality,
since the readily germinable seeds might
produce seedlings of different vigour, some
of which may not result in an established
plant. Therefore, a „Seedling Study‟ was
conducted to evaluate the characteristics
of seedlings grown from seeds produced
by the 5 legume genotypes that had been
grown on the different soil types.
MATERIALS AND METHODS
Germination Study.
Experimental Design. The „Germination
Study‟ was carried out in a growth cabinet
in the School of Marine and Tropical Biology,
James Cook University, Townsville. The
treatments considered 2 factors: the different
legume genotypes (Amar et al., 2012), namely
Stylosanthes, Centrosema, Desmanthus, and
Macroptilium, and the different maternal
backgrounds of the seeds, i.e. seeds of each
of the legumes were collected from the
plants grown on the different soils studied
in the previous Early Growth Study
(Amar et al., 2012). The numbers of seeds
varied between genotypes because of the
variation in numbers of seeds produced.
The limited seed availability made it
impossible to follow the standard procedure
of the International Seed Test (ISTA 1985),
therefore the method of Williams et al.
(1992), who studied germination of small
collections of Acacia seed, was adopted.
The total number of seeds in each treatment
combination (genotype - soil background)
are listed in Table 1.
The 16 treatment combinations
(8 plant genotypes x 2 soil types of
the maternal seed-producing plants)
were arranged in a randomised block
design with 6 replicates resulting in 96
experimental units.
Experimental Procedure. Before germination,
the seeds were put into nylon-bags then
immersed in water at 80oC for 5 minutes,
except for the Stylosanthes scabra cv. Seca
seeds, which were put into the water for 5
2
minutes when the temperature dropped to
75oC. Gilbert and Shaw (1979) found that
Seca seeds achieved their highest germination
percentage (64%) when treated with water
at 75oC.
Seeds were placed in petri-dishes
lined with filter paper following the
international rules for seed testing
guidelines (ISTA 1985), and modified by
putting a layer of 2.5 g vermiculite under 3
layers of filter papers in each petri-dish. The
petri-dishes were then arranged in a
randomised block design with 6 replicates
in a growth cabinet. The growth cabinet
maintained a 12-hour light and dark regime
intermittently every day, with a maximum
temperature of 30 to 32oC, a minimum
temperature of 26 to 28oC, and relative
humidities of 64 to 72% and 74 to 84%, for
light and dark periods, respectively. The
petri-dishes were kept moist for the duration
of the study. The seeds were initially
incubated for 4 weeks due to the very slow
germination of some genotypes, particularly
the Centrosema and Stylosanthes, which
took up to the end of the fourth week to
germinate. The seeds that had not germinated
after 4 weeks were mechanically scarified
by cutting the seed coat with a scalpel at the
opposite end to the embryo. This was done
to facilitate easy germination of hard seeds.
The study was then continued for a further
10 days. By the end of the additional 10
day period, all the seeds that still had not
germinated were considered to be non-viable.
A seed was considered to have
germinated when it developed a radicle more
than 3 mm long (Olff et al., 1994). The
number of germinated seeds was recorded
daily at midday and were removed from the
petri-dishes. At the end of the study period,
seeds were grouped into 3 categories: the
„readily germinable seeds‟ which germinated
within 28 days; the „hard seeds‟ that
germinated after scarification of the seed coat
(within 10 additional days), and the „nonviable seeds‟ that did not germinate by the
end of the experiment, and might comprise
both „dead‟ and „empty‟ seeds according to
(ISTA 1985) definitions. The last two were
not differentiated in this study.
Seed quality was also determined
by the time to 50% germination of the
total number of readily germinable seeds
(T50) using the formula of Coolbear
et al. (1984):
T50 = ti+ {0.5 (N+1) - ni}{ tj - ti} / {nj - ni }
where:
T50 is time to 50% germination,
N is the final total number of readily
germinable seeds, and
„ni‟ and „nj‟ are the number of readily
germinable seeds between two adjacent
counts at times „ti‟ and „tj‟, where “ni<
0.5(N+1)
ISSN : 2407 - 7585
E-ISSN : 2407 - 7593
QUALITY OF SEED PRODUCED BY TROPICAL FORAGE LEGUMES
ON LOW FERTILITY SOILS
Andi L. Amar1), Robert A. Congdon 2), Christopher P. Gardiner3), Ross J. Coventry2)
1)
Faculty of Animal Husbandry and fisheries. University of Tadulako. Palu. 94117 Central Sulawesi. Indonesia.
2)
School of Marine and Tropical Biolog James Cook University. Townsville. Queensland. 4811Australia.
3)
School of Veterinary and Biomedical Science James Cook University. Townsville. Queensland. 4811Australia.
ABSTRACT
Soil fertility can influence seed production and seedling growth. This study compared seed
yields, seed and seedling characteristics of eight forage legumes (Stylosanthesscabra, S. hamata,
Centrosemabrasilianum, C. pascuorum, Desmanthuspubescens, D. Virgatus, Macroptiliumbracteatum
and M. martii) grown on two soil types (red and yellow kandosols), of low fertility, that are
widespread in the Australian tropical rangelands. Seeds of the Desmanthus species germinated most
rapidly and, with M. bracteatum, produced the highest proportion of readily germinable seed. M.
bracteatum seed produced on the Yellow Kandosol germinated significantly faster, as did the S.
hamata from the Red Kandosol than the other species. S. scabra cv. Seca and C. brasilianum
produced more readily germinable seeds from plants grown on the Red Kandosol. C. pascuorum,
D. pubescens and M. bracteatum produced a higher percentage of readily germinable seeds and
fewer hard seeds when grown on the Yellow Kandosol. No significant differences were found in
seedling vigour or biomass allocation from seeds produced on the different soil types.
Key Words : Fertility, forage, legumes, seed, soil, quality.
INTRODUCTION
It is well recognised that plant
development and seed production are
influenced by the surrounding environment.
The environmental conditions during seed
development, along with genetic characteristics
and agronomic techniques have considerable
effect on seed yield and components
of yield through their effect on plant
reproductive physiology (Martiniello, 1998).
The quality of the seed sown, including its
vigor, determines the success of the resulting
plants; while seed quality is reflected in
seedling density and vigor, uniformity of
crop growth, and plants competitiveness, as
well as forage production insome species
(Hampton, 1995). Many publications have
demonstrated that environmental experiences
(e.g. light, nutrient, temperature, water,) of
the maternal plant have influences on
the performaces of the resulting progeny
(Rossiter, 1996; Galloway, 2002; Etterson
and Galloway, 2002; Galloway, 2005;
Luzuriaga, et al., 2006; Galloway and
Etteson, 2007; Uller, 2008; Wolf and Wade,
2009; Mitrovic, et al., 2010).
Nutrient supplies from the soil
in which a plant grows is often found to
be an important factor contributing to
environment-induced maternal effects.
Nutrient status of a plant may affect the
number, size and quality of seeds produced,
as well as the growth and competitive
ability of the plants progeny (Aarssen and
Burton 1990, Sills and Nienhuis 1995).
This is in agreement with Miao et al.
(1991a, 1991b) who showed that the
maternal nutrient environment influences
the competitive ability of progeny in
Plantago spp.; and Galloway (2001)
proved this with a strong effect of maternal
nutrient environment on the germination of
offspringin herbaceous plant Campanula
americana . Though, this is not always
the case. Luzuriaga, et al., (2006) found
1
that added nitrogen to Sinapis arvensis
plants only slightly reduced the germination
rate of its seeds, but not the germination
percentage.
A plant‟s ability to grow under
certain conditions is not enough to indicate
its ability to persist in an ecosystem for
long periods, as is required for a pasture
species. Production of a large number of
seeds is an important factor controlling
plant persistence, as is the seed's capacity
to germinate, to provide new plants for
regeneration. This is particularly important
for an annual species whose persistence
is strongly dependent on both seed production
and viability. Furthermore, establishment
of a good sward is also an important
criterion for pasture production. Early
seedling development plays an important
role in the long-term growth of a plant,
and in the dynamics of plant populations
in the field (Hendrix 1984; Schmitt and
Antonovics 1986). Several studies have
demonstrated the importance of early
emergence in relation to the competitive
ability of the resulting seedlings (Crawley
and Nachapong 1985; Dolan 1984).
Seeds resulting from a previous
study entitled “an Early Growth Study”
(Amar et al. 2012) were tested toinvestigate
the effects of plant genotype and soil type
on the performance of the progeny of the
studied legumes. Although not as strong as
genotype effects, it was found from the
early study that the soil type significantly
affected herbage yield, time to flowering
and the number of seed produced (Amar
et al., 2012). Therefore, it is hypothesised
that edaphic influences on the parent
plants may further affect the viability of
the seeds produced and the performance
of the resulting seedlings. Consequently,
two experiments were conducted here, as a
„Germination Study‟ and a „Seedling Study‟
which are described below.
The „Germination Study‟ compared
seed germination over time, the proportions
of readily germinable seeds, hard seeds,
and non-viable seeds resulting from the 8
legume genotypes grown on the different
soils. The germination criteria, however,
may not be sufficient to judge seed quality,
since the readily germinable seeds might
produce seedlings of different vigour, some
of which may not result in an established
plant. Therefore, a „Seedling Study‟ was
conducted to evaluate the characteristics
of seedlings grown from seeds produced
by the 5 legume genotypes that had been
grown on the different soil types.
MATERIALS AND METHODS
Germination Study.
Experimental Design. The „Germination
Study‟ was carried out in a growth cabinet
in the School of Marine and Tropical Biology,
James Cook University, Townsville. The
treatments considered 2 factors: the different
legume genotypes (Amar et al., 2012), namely
Stylosanthes, Centrosema, Desmanthus, and
Macroptilium, and the different maternal
backgrounds of the seeds, i.e. seeds of each
of the legumes were collected from the
plants grown on the different soils studied
in the previous Early Growth Study
(Amar et al., 2012). The numbers of seeds
varied between genotypes because of the
variation in numbers of seeds produced.
The limited seed availability made it
impossible to follow the standard procedure
of the International Seed Test (ISTA 1985),
therefore the method of Williams et al.
(1992), who studied germination of small
collections of Acacia seed, was adopted.
The total number of seeds in each treatment
combination (genotype - soil background)
are listed in Table 1.
The 16 treatment combinations
(8 plant genotypes x 2 soil types of
the maternal seed-producing plants)
were arranged in a randomised block
design with 6 replicates resulting in 96
experimental units.
Experimental Procedure. Before germination,
the seeds were put into nylon-bags then
immersed in water at 80oC for 5 minutes,
except for the Stylosanthes scabra cv. Seca
seeds, which were put into the water for 5
2
minutes when the temperature dropped to
75oC. Gilbert and Shaw (1979) found that
Seca seeds achieved their highest germination
percentage (64%) when treated with water
at 75oC.
Seeds were placed in petri-dishes
lined with filter paper following the
international rules for seed testing
guidelines (ISTA 1985), and modified by
putting a layer of 2.5 g vermiculite under 3
layers of filter papers in each petri-dish. The
petri-dishes were then arranged in a
randomised block design with 6 replicates
in a growth cabinet. The growth cabinet
maintained a 12-hour light and dark regime
intermittently every day, with a maximum
temperature of 30 to 32oC, a minimum
temperature of 26 to 28oC, and relative
humidities of 64 to 72% and 74 to 84%, for
light and dark periods, respectively. The
petri-dishes were kept moist for the duration
of the study. The seeds were initially
incubated for 4 weeks due to the very slow
germination of some genotypes, particularly
the Centrosema and Stylosanthes, which
took up to the end of the fourth week to
germinate. The seeds that had not germinated
after 4 weeks were mechanically scarified
by cutting the seed coat with a scalpel at the
opposite end to the embryo. This was done
to facilitate easy germination of hard seeds.
The study was then continued for a further
10 days. By the end of the additional 10
day period, all the seeds that still had not
germinated were considered to be non-viable.
A seed was considered to have
germinated when it developed a radicle more
than 3 mm long (Olff et al., 1994). The
number of germinated seeds was recorded
daily at midday and were removed from the
petri-dishes. At the end of the study period,
seeds were grouped into 3 categories: the
„readily germinable seeds‟ which germinated
within 28 days; the „hard seeds‟ that
germinated after scarification of the seed coat
(within 10 additional days), and the „nonviable seeds‟ that did not germinate by the
end of the experiment, and might comprise
both „dead‟ and „empty‟ seeds according to
(ISTA 1985) definitions. The last two were
not differentiated in this study.
Seed quality was also determined
by the time to 50% germination of the
total number of readily germinable seeds
(T50) using the formula of Coolbear
et al. (1984):
T50 = ti+ {0.5 (N+1) - ni}{ tj - ti} / {nj - ni }
where:
T50 is time to 50% germination,
N is the final total number of readily
germinable seeds, and
„ni‟ and „nj‟ are the number of readily
germinable seeds between two adjacent
counts at times „ti‟ and „tj‟, where “ni<
0.5(N+1)