Seed germination responses of Cereus jamacaru DC. ssp.
jamacaru (Cactaceae) to environmental factorspsbi_274 120..128
MARCOS VINICIUS MEIADO,* LARISSA SIMÕES CORRÊA DE ALBUQUERQUE,†
EMERSON ANTÔNIO ROCHA,‡ MARIANA ROJAS-ARÉCHIGA§ and INARA ROBERTA LEAL¶
*Post-Graduate Program in Plant Biology, †Department of Zoology and ¶Department of Botany, Federal University of
Pernambuco, Av. Professor Moraes Rego, Cidade Universitaria, Recife, Pernambuco, Brazil, ‡Department of Biological Science,
State University of Santa Cruz, Rodovia Ilheus-Ibatuba, Km 16, Salobrinho, Ilheus, Bahia, Brazil; and §Department of
Biodiversity Ecology, Ecology Institute, National Autonomous University of Mexico, Ciudad Universitaria, DF 04510 Mexico
Abstract
In the present study, we assessed the seed germination responses of Cereus jamacaru DC.
ssp. jamacaru (Cactaceae) to environmental factors. The seeds were collected from an area
within Caatinga, a semiarid vegetation area located in north-eastern Brazil. We deter-
mined the optimal temperature for germination of C. jamacaru seeds and evaluated the
effect of temperature, light intensity, light quality, water and saline stress on seed ger-
mination. Cereus jamacaru was classified as a positive photoblastic species. Maximum
germination (95.8  2.6%) was found under white light, and seed germination was not
observed in darkness in any of the temperature, water or saline stress treatments. The
optimum temperature for seed germination was 30°C because this temperature favored
most of the parameters evaluated. Seed germinability responded positively to a wide
range of temperatures, but was affected neither by light intensity nor by light quality. A
reduction in water availability and an increase in saline concentration affected ger-
minability and promoted slower, unsynchronized germination. The positive response of
C. jamacaru seed germination to the environmental factors investigated may account for
the abundant occurrence and wide distribution of the species in the Caatinga area.
Keywords: abiotic factors, photoblastism, saline stress, temperature, water stress.
Received 29 July 2009; accepted 27 January 2010
Introduction
The ability of seeds to germinate under different abiotic
factors is of crucial importance for both the survival and
perpetuation of plant species and forest regeneration
(Ceccon et al. 2006). Light, water availability and tempera-
ture are important abiotic factors that determine the ger-
mination of dispersed seeds (Baskin & Baskin 1998). These
factors may be extreme, so their effects can be crucial
in the germination and establishment of plants inhabiting
arid and semiarid environments (Kigel 1995). Thus, the
role of abiotic factors are essential to understand seed
germination responses, and seedling establishment is
important for the maintenance of the genetic diversity of
natural populations (Harper 1977).
In arid and semiarid environments, many studies
that have examined the effect of abiotic factors on seed
germination have used annual species, which possess a
germination pattern considerably different from perennial
species (Kigel 1995). Among the families of perennial
plants that occur in semiarid environments in America,
we can highlight the Cactaceae (Anderson 2001). Over the
past few decades, many studies examining the influence
of abiotic factors on seed germination of Cactaceae have
been conducted (e.g. Nolasco et al. 1996, 1997; Rojas-
Aréchiga et al. 1997, 1998, 2001; De la Barrera & Nobel
2003; Benítez-Rodríguez et al. 2004; Ramírez-Padilla &
Valverde 2005; Flores et al. 2006; Ortega-Baes & Rojas-
Aréchiga 2007; Gurvich et al. 2008). For more examples
and further information, see the review by Rojas-Aréchiga
Correspondence: Marcos Vinicius Meiado
Email: marcos_meiado@yahoo.com.br
Plant Species Biology (2010) 25, 120–128 doi: 10.1111/j.1442-1984.2010.00274.x
© 2010 The Authors
Journal compilation © 2010 The Society for the Study of Species Biology

and Vázquez-Yanes (2000). According to these studies, the
influences of abiotic factors are crucial in Cactaceae
because seedling recruitment depends on several factors,
including the requirements for seed germination and
seedling establishment (Rojas-Aréchiga & Mandujano
2008).
Although the third highest diversity of species in the
Cactaceae family is found in Brazil (Taylor & Zappi 2004),
few studies have been conducted with cacti inhabiting the
Caatinga vegetation, a semiarid ecosystem that character-
izes north-eastern Brazil (Meiado et al. 2008a). Cactaceae
constitutes one of the most important resources for Caat-
inga fauna (Rocha & Agra 2002; Rocha et al. 2007). Thus,
studies that determine the ability of seeds to germinate
under the influence of different abiotic factors may be
important. The aim of our study was to evaluate the seed
germination responses of Cereus jamacaru DC. ssp.
jamacaru (Cactaceae) to some environmental factors, as
well as to contribute to the knowledge of the ecophysiol-
ogy of a species that is widely distributed in Brazilian
ecosystems.
Materials and methods
Species studied and seed collection
The genus Cereus (Hermann) Miller belongs to the tribe
Cereeae and comprises approximately 35 species, distrib-
uted into four subgenus: Cereus, Ebneria, Mirabella and
Oblongicarpi (Anderson 2001). In addition, two subspecies
of Cereus jamacaru DC. are recognized. The subspecies
jamacaru is widely distributed, both naturally and through
human activity, and the subspecies calcirupicola occurs
only in the State of Minas Gerais, Brazil (Anderson 2001).
We assessed the seed germination responses of Cereus
jamacaru DC. ssp. jamacaru, a columnar cactus popularly
known in Brazil as ‘mandacaru’, to some environmental
factors. This species is used as forage, as an ornamental
plant and in popular medicine (Andrade et al. 2006).
Plants are treelike with many erect branches, forming
dense crowns up to 10 m high, with distinct trunks of
up to 60 cm in diameter. Stems are cylindrical, segmented
and greenish blue, up to 15 cm in diameter, with 4–6 ribs.
The flowers are solitary, nocturnal and white. The fruits
are ellipsoidal, between 5 and 12 cm in length, and
7–12 cm in diameter, red, with white pulp (Anderson
2001). Seeds are primarily dispersed by birds and bats
(M. V. Meiado 2007) and secondarily by ants (Leal et al.
2007). We chose this species because it is widely distrib-
uted and occurs in many Brazilian ecosystems, which
ensures that seeds are exposed to different abiotic factors
in environments where they would be able to germinate.
According to Prisco (1966) and Guedes et al. (2009), ‘man-
dacaru’ seeds are light sensitive for germination and a
temperature of 25°C is not adequate for germination and
vigor tests. Moreover, discontinuous hydration of ‘man-
dacaru’ seeds can promote seed germination (Rito et al.
2009).
Ripe fruits of ‘mandacaru’ were collected in April 2007,
at the end of the rainy season, from 20 adult plants growing
within an area of Caatinga, Serra Talhada municipality,
Pernambuco, north-eastern Brazil (7°59’S, 38°19′W). This
region consists of patches of seasonally dry forest and
sclerophyll vegetation (sensu Mooney et al. 1995; Penning-
ton et al. 2000) covering a 730 000 km2 semiarid region
(Sampaio 1995). Variation in the vegetation structure is
conditioned by topography, human disturbance and, most
importantly, by a combination of the average annual rain-
fall and soil attributes (Sampaio 1995; Prado 2003). Rainfall
ranges from 240 to 900 mm per year throughout the Caat-
inga, and soils range from moderately fertile, saline and
shallow to impoverished deep sandy soils, at both land-
scape and regional levels (Sampaio 1995). The vegetation is
predominantly formed by small, woody and herbaceous
deciduous, caducifolious and spiny species, and the Cac-
taceae are one of the most important plant families repre-
sented in this ecosystem (Taylor & Zappi 2004).
Germination tests
We tested the effects of temperature, light intensity, light
quality, water and saline stress on the seed germination of
C. jamacaru ssp. jamacaru. First, seeds were extracted from
the fruits and immediately used for the experiments. For
all treatments, 100 seeds were placed on Petri dishes with
filter paper humidified with the test solutions (5 mL).
There were five dishes per treatment. The dishes were
sealed with transparent masking tape. No solution was
added into the Petri dishes during the assessment period.
Radicle protrusion was considered to be the criteria for
germinated seeds, and was assessed daily over a 50-day
period. In the present study, germination is defined as the
time when the radicle tip emerged  1 mm from the seed
coat.
Temperature treatments
Initially, to verify the optimal temperature for seed germi-
nation of C. jamacaru, we evaluated the effect of tempera-
ture under six constant treatments: 15, 20, 25, 30, 35 and
40°C. In each treatment, the seeds were placed for germi-
nation under white light (12 h photoperiod with a light
intensity of 200 mmol/cm2/s) and in continuous darkness
(simulated with the use of a black polypropylene plastic)
to determine the seed’s photoblastism under each tem-
perature treatment. Germination was investigated every
day for 50 days. Seeds kept in continuous darkness were
counted 50 days after sowing. After this analysis, all the
SEED GERMINATION OF C E R E U S J A M A C A R U 121
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other experiments (i.e. light intensity, light quality, water
and saline stress) were conducted at the optimal tem-
perature for germination, which was determined in this
experiment.
Light treatments
The different levels of light intensity (0, 30, 50, 70 and
100%, where 100% represented the full light treatment
with a light intensity of 200 mmol/cm2/s measured with
a lux meter; LX1010B Digital Lux Meter, Mastech,
Kowloon, Hong Kong) were simulated with the use of a
black polypropylene plastic (0%) and black polypropylene
shade cloth with different mesh sizes (other light intensi-
ties). Light was obtained from cool, daylight fluorescent
and incandescent tubes. We considered 200 mmol/cm2/s
to be 100% of light intensity because that was the
maximum light intensity achieved inside the germination
chamber. Germination was investigated every day over a
50-day period. Seeds kept in continuous darkness were
counted 50 days after sowing.
The influence of light quality was evaluated by submit-
ting the seeds to white, red, far-red and blue light. Red,
far-red and blue lights were obtained from cool, daylight
fluorescent and incandescent tubes with double layers of
red, red-blue and blue cellophane as filters, respectively.
All light quality treatments had the same light intensity
(200 mmol/cm2/s). Seeds were counted every day over a
50-day period, and under the same light quality.
Water stress and saline stress treatments
We used commercial solutions of polyethylene glycol
(PEG) 6000 (Villela et al. 1991) and sodium chloride (Brac-
cini et al. 1996) to simulate water and saline stress, respec-
tively. In both experiments, we evaluated the osmotic
potentials 0.0 (distilled water), -0.2, -0.4, -0.6, -0.8 and
-1.0 MPa, and the dishes were maintained under light
(12 h photoperiod with a light intensity of 200 mmol/
cm2/s) and continuous darkness (simulated with the use
of a black polypropylene plastic). The osmotic potential
was calculated according to Villela et al. (1991) and Brac-
cini et al. (1996), and measured with an osmometer (Mark
3—Osmometer, Fiske Associates, Norwood, USA) at the
beginning of the treatment. No solution was added to the
Petri dishes during the experiment. Germination was
investigated every day over a 50-day period. Seeds kept in
continuous darkness were counted 50 days after sowing.
Statistical analysis
For each seed germination treatment, we calculated ger-
minability (%), mean germination time (t = Sni.ti/Sni,
where ti is the period from the start of the experiment to
the ith observation (day) and ni is the number of seeds
germinated in the time i [not the accumulated number,
but the number corresponding to the ith observation]) and
the synchronization index (E = -Sfi.log2fi, where fi is the
relative frequency of germination [i.e. the proportion of
germinated seeds in an interval]) according to Ranal and
Santana (2006). The germinability data were transformed
to arcsine✓% (Ranal & Santana 2006). Differences in ger-
mination parameters among treatments were tested for
statistical significance using a one-way anova followed
by a Tukey’s honestly significant difference test. Data were
expressed as mean  standard error (SE) values. We
tested the normal distribution of the data and homogene-
ity of the variances using Shapiro–Wilk and Levene tests,
respectively (Zar 1999). All statistical analyses were
carried out using the program STATISTICA 7.0 (P < 0.05).
Results
Temperature treatments
We found significant differences in seed germinabi-
lity among the temperature treatments (F(5,18) = 17.52,
P < 0.0001; Fig. 1). Temperature influenced mean germina-
tion time (F(5,18) = 91.74, P < 0.0001) and the index of seed
germination synchronization (F(5,18) = 9.519, P = 0.0002;
Table 1). The highest percentages of seed germination were
obtained under 25°C (94.0  1.6%) and 30°C (95.8  2.7%),
with no significant difference between these temperatures
(P = 0.8838). Nevertheless, we chose 30°C as the optimal
germination temperature for the species studied because
most of the parameters evaluated in the present study were
favored by this treatment, resulting in the highest ger-
minability (95.8  2.7%) and the lowest mean germination
Fig. 1 Germinability (mean  standard error) of Cereus jamacaru
DC. ssp. jamacaru seeds under six different treatments of constant
temperature (°C) with a 12-h photoperiod. Different letters indi-
cate significant differences at P  0.05 (Tukey’s honestly signifi-
cant difference test). Values are the results 50 days after sowing.
122 M. V. ME IADO E T A L .
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Journal compilation © 2010 The Society for the Study of Species Biology

time (5.5  0.1 days); this last parameter was significantly
different from 25°C (P = 0.0349 and P = 0.0426).
Light treatments
Cereus jamacaru DC. ssp. jamacaru was classified as a posi-
tive photoblastic species. The maximum germination per-
centage (95.8  2.7%) was recorded under white light
(Fig. 2b), and we found no germination under darkness
(Fig. 2a). Light intensity affected all of the parameters
evaluated, except for germinability. We did not observe
seed germination in darkness, and the germinability
of C. jamacaru was similar in all other light treatments
(F(3,12) = 0.89, P = 0.4697; Fig. 2a). In the light intensity
treatments, we found an increase in the mean germina-
tion time (F(3,12) = 4.19, P = 0.0302) and an increase in
the values of the synchronization index (F(3,12) = 6.03,
P = 0.0096; Table 2) as the light intensity was reduced.
These results indicated that a reduction in light intensity
promoted a slower, more unsynchronized germination,
meaning that the seeds germinated more irregularly
within a time interval.
Light quality did not affect C. jamacaru germinability
(F(3,12) = 2.49, P = 0.1101; Fig. 2b), but all other parameters
were influenced by light quality (mean germination
time: F(3,12) = 8.41, P = 0.0009 and synchronization index:
F(3,12) = 5.61, P = 0.0058; Table 2). White light promoted a
faster and more synchronized germination when com-
pared with the other light quality treatments.
Water stress and saline stress treatments
With respect to the experiments carried out under differ-
ent solutions of PEG 6000, we found no germination
under darkness. In addition, we observed low seed ger-
mination at a solution concentration of -0.8 MPa under
white light (Fig. 3a). However, we observed a decrease
in C. jamacaru seed germinability with a reduction in
water availability (F(5,18) = 48.92, P < 0.0001; Fig. 3a), which
affected the mean germination time (F(5,18) = 35.91,
P < 0.0001) and the synchronization index (F(2,9) = 15.82,
P = 0.0011; Table 3). The -0.6 and -0.8 MPa treatments
were removed from the analyses to obtain synchroniza-
tion index data because the high synchrony found in
these treatments was influenced by the low number of
seeds that germinated on the same day (germinability
< 5%).
Seeds submitted to saline stress treatments also showed
a significant reduction in germinability as the saline solu-
tion concentration increased (F(5,18) = 11.18, P = 0.0001;
Fig. 3b), and we found no germinated seeds under dark-
ness. However, there was no significant difference in the
germinability of the seeds submitted to the treatments
Table 1 Mean germination time (days)
and synchronization index of Cereus
jamacaru DC. ssp. jamacaru seeds submitted
to different treatments of constant tem-
perature with a 12-h photoperiod
Constant temperature (°C)
Mean germination
time (days) Synchronization index
15 18.2  0.3 a 1.6  0.2 c
20 10.0  0.4 b 1.9  0.1 b
25 6.6  0.3 c 2.2  0.2 ab
30 5.5  0.1 d 2.4  0.1 a
35 6.4  0.1 c 2.2  0.2 ab
40 9.4  1.0 b 1.5  0.1 c
Values are the results 50 days after sowing. Different letters indicate significant differ-
ences at P  0.05 (Tukey’s honestly significant difference test).
Fig. 2 Germinability (mean  standard error) of Cereus jamacaru
DC. ssp. jamacaru seeds at different light treatments under 30°C
with a 12-h photoperiod. (a) White light intensity and (b) light
quality. No significant differences were found in any light treat-
ment at P  0.05 (Tukey’s honestly significant difference test).
Values are the results 50 days after sowing.
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-0.2, -0.4 and -0.6 MPa when compared with 0.0 MPa
(P = 0.9998, P = 0.9998, and P = 0.7236, respectively). Salin-
ity also affected the other parameters evaluated (mean
germination time: F(5,18) = 83.35, P < 0.0001 and synchroni-
zation index: F(5,18) = 5.61, P = 0.0058; Table 3), and an
increase in the saline concentration promoted slower,
more unsynchronized germination.
Discussion
In the present study, we demonstrated that ‘mandacaru’
seeds are able to germinate under a wide range of envi-
ronmental conditions, characterizing the germination
behavior of species occurring in semiarid ecosystems.
Beyond germinability, other parameters (e.g. mean germi-
nation time and synchronization index of seed germi-
nation) must be considered in the seed germination
responses to environmental factors. These parameters can
contribute substantially to our understanding of seed ger-
mination processes and seedling recruitment in the field,
which are influenced by many abiotic factors.
With respect to temperature, most cacti species respond
positively over a wide temperature interval (Rojas-
Aréchiga and Vázquez-Yanes 2000). For cacti seed germi-
nation, the favorable temperature range is 17–34°C, with
optimal values frequently around 25°C. When we
assessed the effect of temperature on C. jamacaru seed
germinability, the results indicated an optimal tempera-
ture for germination ranging from 25 to 30°C. However,
the mean germination time showed that germination of
the studied species was faster when the seeds were at
30°C. This response might be favorable for germination of
the species, given that even during the rainy season in
Caatinga the temperature of the soil interface may be high
throughout the day. Other columnar cacti, such as Pachyc-
ereus hollianus (Weber) Buxbaum, Cephalocereus chrysacan-
thus (Weber) Britton & Rose, Neobuxbaumia tetetzo var.
tetetzo (Coult.) Backeb. (Rojas-Aréchiga et al. 1998), Steno-
cereus queretaroensis (Weber) Buxbaum (De la Barrera &
Nobel 2003) and Trichocereus terscheckii (Pfeiff.) Britton &
Rose (Ortega-Baes & Rojas-Aréchiga 2007) also germinate
over a wide range of temperatures. According to Rojas-
Aréchiga et al. (1998), the seed germination response to
temperature may be affected by the life form of the cacti,
Table 2 Mean germination time (days) and
synchronization index of Cereus jamacaru
DC. ssp. jamacaru seeds submitted to differ-
ent treatments of light intensity and light
quality at 30°C with a 12-h photoperiod
Light intensity (%)
Mean germination
time (days) Synchronization index
0 – –
30 9.0  1.0 a 3.0  0.2 a
50 7.4  0.8 ab 3.0  0.1 a
70 7.2  0.4 ab 2.7  0.1 ab
100 5.5  0.1 b 2.4  0.1 b
Light quality
Mean germination
time (days) Synchronization index
White 5.5  0.1 c 2.4  0.1 b
Red 7.2  0.9 bc 2.9  0.1 ab
Far-red 9.0  0.7 ab 3.1  0.2 a
Blue 10.8  0.7 a 3.4  0.3 a
Values are the results 50 days after sowing. Different letters indicate significant differ-
ences at P  0.05 (Tukey’s honestly significant difference test). –, no seed germination.
Fig. 3 Germinability (mean  standard error) of Cereus jamacaru
DC. ssp. jamacaru seeds under two stress treatments. (a) Water
stress and (b) saline stress. Different letters indicate significant
differences at P  0.05 (Tukey’s honestly significant difference
test). Values are the results 50 days after sowing.
124 M. V. ME IADO E T A L .
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Journal compilation © 2010 The Society for the Study of Species Biology

although this has not been shown yet. Our studied
species, similar to other columnar cacti, is able to ger-
minate at a higher temperature range than barrel and
globose cacti (Rojas-Aréchiga et al. 1998). The decrease in
germination of C. jamacaru seeds at extreme temperatures
may have an ecological significance because cacti seedling
survival could decrease at these temperatures.
Another response of cacti seed germination that might
be related to cacti life form is the light requirement for
germination (Rojas-Aréchiga et al. 1997). All studies exam-
ining the effects of light on the germination of cacti seeds
have shown two photoblastic responses: positive or light
indifferent (Rojas-Aréchiga & Mandujano 2008). Cereus
jamacaru seeds have been shown to be light sensitive for
germination (Prisco 1966). In that study, Prisco (1966) used
sand as the substrate for germination, sowing the seeds to
a depth of 0.5 cm. This method, although widely used to
evaluate the germination of many species, does not guar-
antee the total absence of light, which promoted a final
germination percentage of 13% after 12 days of evaluation.
In this same study, in a second experiment, seeds of C.
jamacaru were kept in Petri dishes in darkness and the final
percentage of germination was 2%. In our study, all experi-
ments carried out to test the effect of abiotic factors (tem-
perature, water and saline stress) were also assessed in
darkness and there was no germination of any seed after
50 days. Thus, after evaluating the germination of 9000
seeds in darkness under different abiotic factors, our
results allow us to conclude that the columnar cactus
studied here presents a positive photoblastism in accor-
dance with the general pattern found within Cactaceae.
According to Rojas-Aréchiga et al. (1997), barrel and
globose cacti are positive photoblastic, and columnar cacti
can be positive photoblastic or light indifferent, owing to
a maternal effect induced by temperature; the climatic
conditions that prevail during seed development may
influence different light requirements (Rojas-Aréchiga
et al. 1997). Other studies support this results (e.g. barrel
and globose cacti: Benítez-Rodríguez et al. 2004; Flores
et al. 2006; Rebouças & Santos 2007; Gurvich et al. 2008;
Rojas-Aréchiga et al. 2008; and for columnar cacti: Rojas-
Aréchiga et al. 2001; De la Barrera & Nobel 2003; Ramírez-
Padilla & Valverde 2005; Ortega-Baes & Rojas-Aréchiga
2007; Meiado et al. 2008a). Moreover, other studies relate
the requirement for light to the permanence of seeds in a
soil seed bank. If seeds of some cacti species are in places
with low light, these ungerminated seeds may remain
viable in the soil for several months because they require
light to germinate (Bowers 2000; Rojas-Aréchiga & Batis
2001).
In contrast, the response of cacti seed germination to
light quality is apparently not related to the life form or to
the taxon to which the species belongs because seeds of
both life forms might be influenced by light quality
(Alcorn & Kurtz 1959; Nolasco et al. 1996; Rojas-Aréchiga
et al. 1997; Benítez-Rodríguez et al. 2004; Ortega-Baes &
Rojas-Aréchiga 2007; Rebouças & Santos 2007) and
species within the same genus might display different
responses (Benítez-Rodríguez et al. 2004). Light-quality
response appears to be more related to photoblastism
than to life form and taxon because positive photoblastic
species are more influenced by light quality than light-
indifferent species (Rojas-Aréchiga et al. 1997). For
example, an influence of light quality on the seed germi-
Table 3 Mean germination time (days) and
synchronization index of Cereus jamacaru
DC. ssp. jamacaru seeds under different
concentrations of PEG 6000 and NaCl solu-
tions at 30°C with a 12-h photoperiod
PEG 6000 solution (MPa)
Mean germination
time (days) Synchronization index
0.0 5.5  0.1 d 2.4  0.1 b
-0.2 14.6  1.3 c 4.0  0.1 a
-0.4 17.8  1.6 bc 4.7  0.1 a
-0.6 20.5  1.8 ab < 5%
-0.8 26.0  0.6 a < 5%
-1.0 – –
NaCl solution (MPa)
Mean germination
time (days) Synchronization index
0.0 5.5  0.1 d 2.4  0.1 c
-0.2 7.4  1.1 d 3.9  0.1 b
-0.4 15.3  1.1 c 4.5  0.1 b
-0.6 20.3  1.4 b 4.7  0.1 b
-0.8 26.7  1.2 a 4.6  0.1 b
-1.0 29.1  1.0 a 5.5  0.2 a
Values are the results 50 days after sowing. Different letters indicate significant differ-
ences at P  0.05 (Tukey’s honestly significant difference test). < 5%, germinability < 5%; –,
no seed germination.
SEED GERMINATION OF C E R E U S J A M A C A R U 125
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nation of positive photoblastic species was found in cacti
of different life forms, such as T. terscheckii (Ortega-Baes &
Rojas-Aréchiga 2007) and Melocactus conoideus Buining &
Brederoo (Rebouças & Santos 2007), columnar and
globose cacti, respectively. In the present study, some ger-
mination parameters (i.e. mean germination time and syn-
chronization index of germination) of the photoblastic
seeds of C. jamacaru ssp. jamacaru were affected by light
quality. However, the germination of some positive pho-
toblastic species of Mammillaria was not influenced by
light quality (Benítez-Rodríguez et al. 2004). Thus, the ger-
mination of light-sensitive cacti might or might not be
influenced by light quality, so a general conclusion cannot
be drawn yet. Finally, for light-indifferent species, such as
Pachycereus pringlei (S. Wats.) Britton & Rose (Nolasco
et al. 1996), P. hollianus, C. chrysacanthus and N. tetetzo
(Rojas-Aréchiga et al. 1997) seed germination was not
influenced by light quality.
Aside from light and temperature, other environmen-
tal factors, such as water availability, can affect seed ger-
mination in arid and semiarid ecosystems (Kigel 1995).
According to Kigel (1995), several desert species are able
to germinate at relatively low soil water potential;
however, germinability decreases with a reduction in
water availability. Indeed, even though germination was
not observed in the -1.0 MPa treatment, we did observe
seed germination at low water potential, as well as a sig-
nificant reduction in seed germinability with a decrease
in water availability. Similar results were found by
Ramírez-Padilla and Valverde (2005), where a -0.4 MPa
treatment caused a significant reduction in seed ger-
minability in three Neobuxbaumia species. Nevertheless,
germination of the columnar cactus S. queretaroensis
occurs at more negative potentials (-1.0 MPa) when com-
pared with germination of the studied species (De la
Barrera & Nobel 2003). This trait might be related to the
high level of rain in Caatinga, which, despite being
restricted to a short period over the year (only a few
days), it might reach values of up to 900 mm in some
areas of the ecosystem (Sampaio 1995) and keep the soil
humidity for long enough to complete the germination
process. Thus, species such as ‘mandacaru’ that exhibit a
faster and more synchronized germination may be
favored in these areas. In contrast, the germinability of
some cacti species, such as N. tetetzo var. tetetzo and P.
hollianus, increases with a reduction in water availability
in the soil (Flores & Briones 2001). According to Flores
and Briones (2001), the relationship among germination
patterns and water availability highlights an important
adaptation of species that germinate in arid and semiarid
ecosystems, and these species would have an advantage
in those environments.
Moreover, the study site presents high soil salinity
caused by the high evaporation of soil water and bad soil
drainage, resulting in an accumulation of salt in the soil
(Fassbender & Bornemisza 1987; Mascarenhas et al. 2005).
However, there are no studies detailing soil salinity
values at the study site; it is possible that soil salinity
could reach levels below -0.8 MPa. Yet, the high percent-
age of seeds that germinated under the saline stress treat-
ments (54.5  15.4% at -0.8 MPa) may indicate that C.
jamacaru is able to germinate in soils with high salinity,
and may be considered to be a halotolerant species. Seed-
ling establishment and adult halotolerance have yet to be
determined. Halotolerance has also been observed in
other columnar cacti, such as P. pringlei (Nolasco et al.
1996). In contrast, seed germination of Ferocactus peninsu-
lae (F.A.C. Weber) Britton & Rose, a barrel cactus, was
inhibited in saline concentrations below -0.2 MPa
(Romero-Schmidt et al. 1992).
In conclusion, seed germination responses may have a
direct impact on the distribution and abundance of plant
species (Valverde et al. 2004). Thus, the abundant occur-
rence and wide distribution of C. jamacaru in the Caatinga
might be related to positive seed germination responses
under a wide range of environmental factors. Neverthe-
less, seedlings of this species are not often found in this
ecosystem (Meiado et al. 2008b). This absence might be
related to one or more specific requirements for seedling
survival and recruitment of new individuals in the popu-
lation or to high seedling predation that must be further
investigated.
Hence, the high fruit production per individual, the
high seed production per fruit, and the high seed germi-
nation capacity under the influence of different environ-
mental factors should compensate for the low level of
recruitment and should favor the occurrence and the wide
distribution of the species in the study site.
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